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BULLETIN
of the
BRITISH
ORNITHOLOGISTS' CLUB
C. W. BENSON
Volume 93
1973
price: three pounds fifty pence (per volume)
PREFACE
There has been one change of importance in the
present volume. As authorised at the 1972 Annual
General Meeting {Bull. Brit. Orn. CI. 92 : 69-70), in
order to offset a further increase in printing and distri-
bution charges, publication of the Bulletin has been
reduced from six to four numbers in the year, al-
though the total number of pages has been maintained.
Those who have been kind enough to advise on
individual papers include Mr. J. H. R. Boswall, Dr.
L. H. Brown, Sir Hugh Elliott, Mr. D. Goodwin,
Mrs. B. P. Hall, Dr. A. R. Phillips, the Rev. Dr. W.
Serle, Prof. W. H. Thorpe, Mr. R. Wagstaffe and Dr.
C. A. Wright. Again I am most grateful to Mr. K. E.
Wiltsher, Manager of the Caxton and Holmesdale
Press, assisted by Mr. P. G. Ball, for the regular pro-
duction of the Bulletin. Also again I wish to thank
Mrs. C. W. Benson, who has spent much time on
proof-reading, and Mrs. M. Hawksley on the com-
pilation of the Index to Scientific Names.
Appreciation of the assistance rendered by Dr.
P. J. K. Burton in the finalisation of the Bulletin for
December 1972 was expressed in the preface for 1972.
The opportunity is now taken to thank him for the
still more important part which he played in the
production of the number for March 1973. Before my
departure for Madagascar it was possible only to hand
over edited typescript the equivalent of rather less
than half of that number. All further editorial work
was carried out by Dr. Burton, and I was only able to
resume duty with the number for June.
C. W. BENSON
LIST OF MEMBERS AS AT 31st DECEMBER, 1973
(Compiled by the Hon. Secretary)
Amended as follows :
New Members
Armistead, H. T., m.s., 39 Benezet Street, Philadelphia, Pennsylvania 191 18, U.S.A.
Carpenter, C. P., 162 Wickham Way, Beckenham, Kent BR3 3AS.
Chatfield, D. G. P., 3 Cyncoed Close, Dunvant, Swansea, Glamorgan SA2 7RS.
Cole, L. R., Commcen, R.A.F. Bawtry, near Doncaster, Yorkshire DN10 6JS.
Davison, G. W. H., 8 Eastwick Road, Walton-on-Thames, Surrey.
El kins, D. A., Little Tickeridge, Kingscote, East Grinstead, Sussex.
Farrand, J., Jnr., m.s., Ornithology Dept., American Museum of Natural History,
Central Park West at 79th St., New York, N.Y. 10024, U.S.A.
Forrester, R. W., a.i.b., 29 Crandleyhill Road, Prestwick, Ayrshire KA9 2BE.
Ginn, H. B., m.a., c/o British Trust for Ornithology, Beech Grove, Tring, Hertfordshire.
Harvey, W. G., m.a., 17 Barton Road, Canterbury, Kent.
Holmes, D. A., b.sc, The Old Rectory, Sudborough, Kettering, Northamptonshire.
Heigham, J. B., m.a., African Timber and Plywood, P.M.B. 4001, Sapele, Midwest State,
Nigeria.
Jobling, J. A., 28 Fairweather House, Parkhurst Road, London N7 oNS.
King, D. G., Fish and Wildlife Branch, 1600 3rd Avenue, Prince George, British Columbia
VZL-3G6, Canada.
Knox, A. G., 9 Urquhart Street, Aberdeen AB2 iPL.
Lawrence, D. W., 62 The Boulevard, West Worthing, Sussex BN13 iLA.
McAndrew, R. T., b.sc, 103 Granville Avenue, Hartlepool, Co. Durham TS26 8NA.
Mackrill, E. J., Mill Lane, Welton-le-Marsh, Spilsby, Lincolnshire.
Medway, Lord, ph.d, Great Glemham House, Saxmundham, Suffolk IP17 iLP.
Meriwani, Y. N., Arasat Al-Hindiyah 389, Baghdad, Iraq.
Neighbour, Lieut.-Colonel J. R., 9 Coldstream Gardens, West Hill Road, London S.W. 18.
Oliver, P. J., 53 Ember Farm Way, East Molesey, Surrey.
Perrin, Dr. M. R., ph.d., b.sc, 12 McGregor Crescent, Pinawa, Manitoba, ROE iCO,
Canada.
Ridland, J. R. J., Troswickness, Dunrossness, Shetland.
Smith, G. A., m.r.cv.s., 158 Broadway, Peterborough PEi 4DC.
Sugg, M. St. J., m.a., 5 The Limes, Hitchin, Hertfordshire SG5 2 AY.
Taylor, R. C, b.sc, Biology Dept., Liverpool Polytechnic, Byrom Street, Liverpool
L 3 3 AF.
Templeton, R. K., The Old Manse, Doune, Perthshire.
Wheeler, C. E., 3 Woodhurst Close, Cuxton, Rochester, Kent.
Zonfrillo, B., 28 Brodie Road, Glasgow G21 3SB.
Resignations
Glenister, A. G.
Markus, M.
Medhurst, H. P.
Deaths
The Committee regrets to record the deaths of the following members : —
Mr. R. P. Borrett
Mr. K. D. Smith
LIST OF AUTHORS AND CONTENTS
Annual General Meeting
Ash, J. S.
Six species of birds new to Ethiopia
Bayliss-Smith, T. P.
A recent immigrant to Ontong Java atoll, Solomon Islands
Beesley, J. S. S.
The breeding seasons of birds in the Arusha National Perk, Tanzania
Benson, C. W. and Irwin, M. P. Stuart
P achy coccyx audeberti: some addenda
BOSWALL, JEFFERY
Supplementary Notes on the Birds of Point Tombo, Argentina
The Nesting of Feral Pigeons Columba livia in Trees
A Discography of Bird Sound from the Oriental Zoogeographical Region
Britton, P. L.
Seasonal movements of the black cuckoo-shrikes Campephaga phoenicea and C,
flava, especially in eastern Africa
Brooke, R. K. and Kemp, A. C.
Specimen data on Bucotvhs leadbeateri
Brown, L. H.
An address on British birds of prey (with precis)
Brown, L. H. and B. E.
The relative numbers of migrant and resident rollers in eastern Kenya
Burton, P. J. K.
Composite nest of Short-crested Flycatcher Myiarchus ferox
Non-passerine bird weights from Panama and Colombia
Button, E. L.
The Common Snipe Gallinago gallinago in Zambia
Chapin, Ruth Trimble
Observations on Bradypterus carpalis and Bradypterus graueri
Chappuis, C. and Erard, C.
A new race of Pectoral-patch Cisticola from Cameroun
Clancey, P. A.
The Status and Characters of the Races of the Red-backed Shrike Wintering
in the South African Sub-Region
Colebrook-Robjent, J. F. R.
The breeding of the Madagascar Banded Kestrel
Colebrook-Rob)ent, J. F. R. and Osborne, T. O.
Nesting of Verreaux's Eagle Aquila verreauxi in Zambia
Collins, Charles T.
The natal pterylosis of the Swallow T -tanager
Committee
Report for 1972
Cooper, John E.
Blood parasites from a Red-chested Owlet Glaucidiuw tephronotum
Dasgupta, J. M.
See Mukherjee
Dean, W. R. J.
Notes on a Lanner with malformed bill, and on hornbills feeding on oil palm
fruits
De Naurois, R.
See Erard
Dickerman, Robert W.
A review of the Boat-billed Heron Cochlearius cocblearius
Elgood, J. H.
Rufous phase Senegal Coucals in southwest Nigeria: an illustration of
Gloger's Rule . .
49-
-52
3-6
5 2
-53
IC
-20
160-
161
33
38
170-
-36
-39
172
41
-47
89
-92
137
126-
130
114-
116-
115
118
174
167-
170
143-
144
92
-96
108-
in
166
155-
157
25-26
55
111-114
i73
Elliott, Hugh F. I.
See Thomas
Erard, C. and De Naurois, R.
A new race of Thekla Lark in Bale, Ethiopia . . . . . . . . 1 41-142
Erard, C. and Jarry, G.
A new race of Thekla Lark in Harrar, Ethiopia . . . . . . . . 139-140
See also Chappuis
Farkas, T.
The biology and a new subspecies of Monticola sharpei .. .. .. 145-155
Farrand, John and Olson, Storrs L.
The correct spelling of Scopoli's specific name for the Malaysian Crested
Wood Partridge (Rollulus) . . .. .. .. .. .. .. 5 3—54
Feare, C. J.
Numenius minutus, Falco subbuteo and Caprimulgus europaeus in the Seychelles 99-101
Field, G. D.
Ortolan and Blue Rock Thrush in Sierra Leone . . . . . . . . 81-82
Subalpine and Grasshopper Warblers in Sierra Leone .. .. . .101-103
Fjeldsa, Jon
Possible Female Hybrids between Bucepbala islandica and clangida . . . . 6-9
Frost, P. G. H.
See Siegfried
Fry, C. H.
The juvenile plumage of Pseudbirimdo griseopyga and identity of "Vom
Swallows"
Goodwin, Derek
Some calls and behaviour patterns of the Plumbeous and Rufous Pigeons . .
Harrison, C. J. O.
The humerus of Ichthyornis as a taxonomically isolating character
Harrisson, Tom
An illustrated address on the birds and bird cult of Easter Island
Holyoak, D. T.
Notes on the birds of Rangiroa, Tuamotu Archipelago, and the surrounding
ocean
Ingram, Collingwood
Exhibition of down feathers of an Eider
Irwin, M. P. Stuart
See Benson
Jackson, H. D.
The Cape Eagle Owl Bubo capensis in Mocambique . . . . . . . . 10
Jarry, G.
See Erard
Jehl, J. R., Rumboll, M. A. E. and Winter, J. P.
Winter bird populations of Golfo San Jose, Argentina . . . . . . 56-63
Keith, Stuart
The voice of Sarotbrura insularis with further notes on members of the genus 130-136
Kemp, A. C.
See Brooke
King, David G.
The birds of the Shira Plateau and west slope of Kibo, Kilimanjaro . . 64-71
Larmuth, John
Migration of Motacilla alba alba . . . . . . . . . . . . 97~9 8
Mees, G. F.
Once more : the identity and authorship of Treron griseicauda . .
Mountfort, Guy
An illustrated address on a visit to India, Pakistan and Nepal
Mukherjee, A. K. and Dasgupta, J. M.
On the taxonomic status of the genus Sauropatis Cabanis & Heine
Alcedinidae)
Nichols, Holly A. J. and Nichols, Thomas Duncan
St. Vincent Parrot : plumage polymorphism, juvenile plumage and nidification
138-
-139
103-
-108
123-
-126
49
26-32
97
. . 119-120
49
(family
.. 79-81
Olson, Storrs L.
See Farrand, Ripley
Osborne, T. O.
See CoLEBROOK-ROBJENT
Parslow, J. L. F.
Organochlorine insecticide residues and food remains in a Bald Ibis Geronti-
cus eremita chick from Birecik, Turkey
Phillips, Allan R.
On the supposed genus Petrochelidon
Prescott, Kenneth W.
First report of Pitta e. erythrogaster from Leyte
A small collection of birds from Muara Island, north Borneo
Prigogine, A.
The migratory movements of the Pygmy Kingfisher Geyx picta natalensis ii
the Republic of Zaire ....
Ripley, S. Dillon and Olson, Storrs L.
Re-identification of Rallus pectoralis deignani
Rowan, M. K.
An illustrated address on African brood parasites
Rumboll, M. A. E.
See Jehl
Schifter, Herbert
A specimen of Coua delalandei (Temminck) (Cuculidae) in the Naturhistori
sches Museum, Vienna (Austria)
Schuz, E.
Some remarks on the Lammergeyer Gypaetus barbatus in East Africa
Short, Lester L.
Remarks on the status of Campethera " ' scriptoricauda\ and related species
Sibley, Charles G.
The Relationships of Picathartes
The Relationships of the Swallow-tanager Tersina viridis
Siegfried, W. Roy
Wing moult of Ruddy Ducks in Manitoba
Siegfried, W. Roy and Frost, P. G. H.
Regular Occurrence of Porphyrula martinica in South Africa
Spencer, Robert
An illustrated address on feeding by Siskins in gardens
Thomas, D. Keith and Elliott, Hugh F. I.
Nesting of the Roseate Tern {Sterna dougallii) near Dar es Salaam
Van Someren, G. R. Cunningham
Tringa hypoleucos Linnaeus breeding in East Africa
Walters, Michael
The eggs of Carpococcyx renauldi
White, C. M. N.
Diomedea cauta in South African waters
Migrant Pernis apivorus in the Indonesian Archipelago
Coracina papuensis in Indonesia
Night Herons in Wallacea
Winter, J. P.
See Jehl
163-166
32-33
157-159
115
1
2-3
. 161-
163
72
-74
23
75
- 2 5
-79
. 98
-99
. 36
-38
97
21
-23
39
-40
174
• 159-
• 175-
56
116
160
176
INDEX TO SCIENTIFIC NAMES
(Compiled by the Editor with the assistance of Mrs. M. Hawksley)
All generic and specific names (of birds only) are indexed. Only new subspecific names are
included, and are indexed under the generic and the specific names.
abbotti, Trichastoma 172
abingoni, Campethera 16, 74
abyssinica, Coracias 130
— Hirundo 17
abyssinicus, Bucorvus 89-90
Acanthis flammea 77
Acrocephalus arundinaceus 31
— atypha 31-32
— baeticatus 5-6
— caffra 31
— scirpaceus 102-103
Actitis macularia 117
Actophilornis africanus 12-13
acutipennis, Chordeiles 118
adustus, Alseonax 15-16
Aegithina tiphia 158-159
— viridissima 158
aegyptiacus, Alopochen 12
aenea, Chloroceryle 1 1 8
aeneus, Dicrurus 172
aequatorialis, Apus 65, 70
aequinoctialis, Procellaria 58-59
aethereus, Phaethon 28
aethiopicus, Laniarius 16-19
Aethopyga siparaja 159
afer, Nilaus 16
— Parus 153
affine, Malacopteron 171
affinis, Sarothrura 70, 133
africana, Mirafra 14-15
africanus, Actophilornis 12-13
— Bubo 26
— Buphagus 16
Agapornis fischeri 17
aguimp, Motacilla 16
alba, Calidris 58, 63
— Chionis 35, 58, 63
— Gygis 30
— Motacilla 97-98
— Pterodroma 27
— Tyto 26
albicollis, Corvus 66
— Nyctidromus 118
albifrons, Amblyospiza 14
— Petrochelidon 20
albigularis, Laterallus 117
— Nesofregetta 28
albiventer, Phalacrocorax 34-35, 58, 60
albiventris, Halcyon 15
albospecularis, Copsychus 152, 154
alboterminatus, Tockus 15, 55
albus, Casmerodius 58, 61
Alcedo senegalensis 79
Alopochen aegyptiacus 12
Alseonax adustus 15-16
Amazona guildingii 120-123
— ochrocephala 1 1 8
amazona, Chloroceryle 118
Amblyospiza albifrons 14
americana, Chloroceryle 118
americanus, Daptrius 117
amethystina, Chalcomitra 17-1!
amicta, Nyctiornis 171
Anas capensis 12
— erythrorhynchus 12
— fiavirostris 58, 61
— georgica 58, 61
— punctata 12
— sparsa 12, 70
Andropadus importunus 4
angolensis, Monticola 154
Anhinga anhinga 117
anhinga, Anhinga 117
ani, Crotophaga 118
Anorrhinus galeritus 172
Anous stolidus 29-30
— tenuirostris 29-30
anthracinus, Buteogallus 117
Anthracoceros malayanus 171
Anthreptes collaris 17-19
— malacensis 159
Anthus similis 14, 18-19, 8l
— trivialis 81
Apalis flavida 16-19
apivorus, Pernis 116
Aplonis feadensis 53
Apus aequatorialis 65, 70
— caffer 17-18
— horus 17-18
— melba 65, 71
— myoptilus 70
— niansae 17-18
Aquila fasciata 10
— rapax 13
— verreauxi 70, 166
Aramides cajanea 117
Archaeopteryx 125-126
ardosiaceus, Falco 108-109
argus, Argusianus 172
Argusianus argus 172
ariel, Fregata 28
Arizelocichla milanjensis 15
— nigriceps 15
armatus, Hoplopterus 14-15
arminjoniana, Pterodroma 27
aromatica, Treron 119
arquatrix, Columba 17, 19
arundinaceus, Acrocephalus 31
Asio capensis 26
astrild, Estrilda 14-15
Ateleodacnis 78
ater, Haematopus 58, 62
atratus, Coragyps 117
atriceps, Phalacrocorax 35, 58, 61
atypha, Acrocephalus 31-32
audeberti, Pachycoccyx 1 60-1 61
aura, Cathartes 117
aurantiigula, Macronyx 14-15, 19, 70
auratus, Hartlaubius 153
aurea, Jacamerops 118
australis, Megalaima 171
— Oxyura 98
— Tchagra 16, 18-19
— Treron 17
avosetta, Recurvirostra 40
ayresii, Sarothrura 132
Aythya erythrophthalma 12
baboecala, Bradypterus 168-169
badia, Ducula 172
baeticatus, Acrocephalus 5-6
bairdii, Calidris 58, 63
Balearica regulorum 12-13
barbatus, Gypaetus 65, 68, 161-162
— Pycnonotus 4, 16-19
Baryphthengus martii 118
Batis molitor 16-19
belcheri, Pachyptila 59
Belonopterus chilensis 117
bennettii, Campethera 72-74
bensoni, Monticola 145
bergii, Sterna 29
bernsteinii, Ptilinopus 120
biarmicus, Falco 55
bicalcaratus, Francolinus 144
bicolor, Spreo 153
— Trichastoma 24
boehmi, Sarothrura 131
borin, Sylvia 102-103
bougainvillii, Phalacrocorax 34-35
brachyptera, Cisticola 14, 144
Bradypterus baboecala 168-169
— carpalis 167-169
— grandis 169
— graueri 167-169
— yokanae 167
bres, Criniger 171
brevicaudata, Camaroptera 16
brodiei, Glaucidium 172
brunnescens, Cisticola 64-65, 69-70, 143-
144
Bubo africanus 26
— capensis 10, 69
— lacteus 13, 26
Buccanodon leucotis 15
Bucephala clangula 6-9
— islandica 6-9
Buceros nagtglasii 120
— pulchrirostris 11 9-1 20
— rhinoceros 172
Bucorvus abyssinicus 89-90
— leadbeateri 89-91
bullockoides, Melittophagus 16
Bulweria 28
Buphagus africanus 16
— erythrorhynchus 16
Burhinus capensis 14
burtoni, Serinus 15
Busarellus nigricollis 117
Butastur rufipennis 129
Buteo magnirostris 117
— oreophilus 13, 65, 68
— platypterus 117
— ■ rufofuscus 13, 19, 65
Buteogallus anthracinus 117
— subtilis 117
— urubitinga 117
Butorides virescens 117
Bycanistes sharpii 55
cachinnans, Herpetotheres 117
caerulea, Florida 117
caffer, Apus 17-18
caffra, Acrocephalus 31
cajanea, Aramides 117
Calamocichla newtoni 133
caledonicus, Nycticorax 175
Calidris alba 58, 63
— bairdii 58, 63
— canutus 58, 63
— fuscicollis 58, 63
— melanotos 3, 6
Caloenas nicobarica 53
Calospiza 76
calvus, Geronticus 163
Camaroptera brevicaudata 16
Campephaga flava 41-43, 46-47
— petiti 41
— phoenicea 41, 44-47
— quiscalina 41
— sulphurata 16
Campephilus melanoleucos 118
Campethera abingoni 16, 74
— ■ bennettii 72-74
— - nubica 16, 72-74
— - punctuligera 72-74
— scriptoricauda 72-74
cancellatus, Prosobonia 28
canicapilla, Nigrita 15
canicollis, Serinus 67
cantans, Cisticola 14, 18-19
cantillans, Sylvia 101
canutus, Calidris 58, 63
capense, Daption 58
capensis, Anas 12
— Asio 26
— Bubo 10, 69
— Burhinus 14
— Euplectes 14
— Pelargopsis 158
— Zonotrichia 77
capicola, Streptopelia 17
Caprimulgus europaeus 99-101
— fossii 14, 19
— fraenatus 14, 19
— - macrurus 172
— poliocephalus 14, 19, 71
cardinalis, Eos 52-53
Carduelis citrinelloides 17-19
carpalis, Bradypterus 167-169
Carpococcyx renauldi 174
Casmerodius albus 58, 61
cassinii, Leptotila 117
castaneiceps, Ploceus 14-15
Catharacta skua 35, 58, 63
Cathartes aura 117
Catoptrophorus semipalmatus 117
caudata, Coracias 127-129
cauta, Diomedia 56
cayana, Piaya 118
cayennensis, Columba 103
Cecropis 20
celebensis, Pernis 116
Celeus loricatus 1 1 8
Centropus epomidis 173
— senegalensis 173
— superciliosus 14
Centurus pucherani 1 1 8
— rubricapillus 118
Cercomela sordida 64-66, 70
cerulea, Procelsterna 29
Cer yle torquata 1 1 8
Ceyx picta 82, 84-85
Chalcomitra amethystina 17-18
— senegalensis 17-18
Charadrius falklandicus 35, 58, 62
— pecuarius 14
— tricollaris 14
— wilsonia 117
cheela, Spilomis 171
chelicuti, Halcyon 16
chilensis, Belonopterus 117
— Phoenicopterus 35, 58, 61
— Vanellus 35, 58
chimachima, Milvago 117
chiniana, Cisticola 14
Chionis alba 35, 58, 63
Chlidonias niger 1 1 7
Chloephaga picta 58, 61
— rudiceps 61
chloris, Halcyon 80, 158
Chlorocer yle aenea 1 1 8
— amazona 118
— americana 118
— inda 118
Chlorocichla flavicollis 4, 6
Chloropeta natalensis 14
Chlorophoneus sulfureopectus 16, 18-19
chloropus, Gallinula 12
Chordeiles acutipennis 118
chrysaea, Stachyris 172
Ciccaba woodfordii 26
cinclorhynchus, Monticola 149
cinereiceps, Ortalis 117
Cinnyricinclus leucogaster 1 60-1 61
Cinnyris mediocris 15
— venustus 17-19
cirlus, Emberiza 77
cirrhatus, Spizaetus 172
Cisticola 103
— brachyptera 14, 144
— brunnescens 64-65, 69-70, 143-
144
Cisticola brunnescens mbangensis,
subsp. nov. 144
Cisticola cantans 14, 18-19
— chiniana 14
— galactotes 12-13
— hunteri 64, 66, 70
— natalensis 144
— woosnami 14, 19
citrinelloides, Carduelis 17-19
clangula, Bucephala 6-9
clara, Motacilla 12-13
Claravis pretiosa 117
Coccopygia melanotis 14-15
Cochlearius cochlearius 111-113
cochlearius, Cochlearius m-113
Coereba 78
— flaveola 77
Colius striatus 16, 18-19
Coliuspasser laticauda 14
collaris, Anthreptes 17-19
— Lanius 16-19, 7°
Collocalia maxima 171
collurio, Lanius 92-96
Columba arquatrix 17, 19
— cayennensis 103
— cristata 53
— leucocephala 107
— livia 38-39, 107
— maculosa 107
— nigrirostris 117
— picazuro 105, 107
— plumbea 103-107
— subvinacea 104
Columbina minuta 117
— talpacoti 117
concolor, Falco 101
Conirostrum 78
Copsychus albospecularis 152, 154
— malabaricus 1 71-172
— saularis 172
Coracias abyssinica 130
— caudata 127-129
— garrulus 126-130
— naevia 127-130
Coracina papuensis 1 5 9
Coracopsis vasa 153
Coragyps atratus 1 1 7
coralensis, Ptilinopus 30-31
coromanda, Halcyon 80
coronatus, Stephanibyx 14
Corvus albicollis 66
— enca 172
Cossypha imerina 145
— natalensis 43
— semirufa 15
— sharpei 147
Coturnix coturnix 68
coturnix, Coturnix 68
Coua delalandei 2-3
— serriana 3
crestatus, Eudyptes 34
Criniger bres 171
— phaeocephalus 171
cristata, Columba 53
— Fulica 12
croceus, Macronyx 144
Crotophaga ani 118
— major 118
— sulcirostris 118
cubla, Dryoscopus 15
cucullatus, Orthotomus 172
— Spermestes 14
Cuculus micropterus 171
— sparverioides 171
Cuncuma vocifer 13
cuneata, Geopelia 105
cuprea, Nectarinia 144
curvirostra, Treron 158
cuvieri, Dryolimnas 133
cyanea, Passerina 77
Cyanerpes cyaneus 78
cyaneus, Cyanerpes 78
cyanomelas, Rhinopomastus 16
Cyanomitra olivacea 15-16
Cygnus melanocoryphus 35
Daption capense 58
Daptrius americanus 117
delalandei, Coua 2-3
demersus, Spheniscus 34
Dendroica pensylvanica 77
— petechia 77
Dendropicos fuscescens 16
Dicaeum trigonostigma 159
dickinsoni, Falco 108-109
Dicrurus aeneus 172
— paradiseus 171
Diomedia cauta 56
— melanophris 34, 58-59
Dioptrornis fischeri 16, 19
dispar, Ichthyornis 124-125
Diuca diuca 36, 77
diuca, Diuca 36, 77
dominica, Pluvialis 28
dominicanus, Larus 33, 36, 58-59, 63
dominicus, Podiceps 117
dougalii, Sterna 21
Dryocopus javensis 171
Dryolimnas cuvieri 133
Dryoscopus cubla 15
Ducula badia 172
— pacifica 53
dumetaria, Upucerthia 36
dupetithouarsii, Ptilinopus 31
duvauceli, Harpactes 172
ecaudatus, Terathopius 13
Egretta sacra 28
Elanoides forficatus 117
Electron platyrhynchum 118
elegans, Sarothrura 134-135
eleonorae, Falco 101
Emberiza cirlus 77
— ■ hortulana 81
— tahapisi 81
enca, Corvus 172
Eos cardinalis 52-53
Ephippiorhynchus senegalensis 12-13
episcopus, Thraupis 77
epomidis, Centropus 173
eremita, Geronticus 163
Ereunetes mauri 117
erythrocephalus, Garrulax 172
erythrogaster, Pitta 32-33
erythromelas, Piranga 76
erythrophthalma, Aythya 12
erythrorhynchus, Anas 12
— Buphagus 16
Estrilda astrild 14-15
Eucometis penicillata 77
Eudynamis taitensis 52
Eudyptes crestatus 34
Eupetes 24
Euphonia luteicapilla 77
Euplectes capensis 14
— flammiceps 161
— hordeaceus 161
europaeus, Caprimulgus 99-101
Eurostopodus temminckii 171
eurycricotus, Zosterops 15
eurygnatha, Sterna 36
Eurystomus glaucurus 16-17, I 3°
Falco ardosiaceus 108-109
— biarmicus 55
— concolor 101
— dickinsoni 108-109
— ■ eleonorae 101
— femoralis 58, 61
— newtoni 108-109
— peregrinus 13, 38, 61, 108
— subbuteo 99-101, 127
— zoniventris 108-111
falklandicus, Charadrius 35, 58, 62
famosa, Nectarinia 153
fasciata, Aquila 10
feadensis, Aplonis 53
femoralis, Falco 58, 61
ferox, Myiarchus 114
fischeri, Agapornis 17
— Dioptrornis 16, 19
flammea, Acanthis 77
flammiceps, Euplectes 161
flava, Campephaga 41-43, 46-47
flaveola, Coereba 77
flavicollis, Chlorocichla 4, 6
flavida, Apalis 16-19
flavipes, Tringa 117
flavirostra, Limnocorax 12
flavirostris, Anas 58, 61
— Tockus 55
flavivertex, Serinus 17-18
flavus, Xanthopsar 77
Florida caerulea 117
fluviatilis, Locustella 4, 6
forficatus, Elanoides 117
formosus, Ptilinopus 120
fortis, Geospiza 77
fossii, Caprimulgus 14, 19
fraenatus, Caprimulgus 14, 19
Francolinus bicalcaratus 144
— hildebrandti 14
— psilolaemus 70
— shelleyi 70
— squamatus 14
Fregata ariel 28
— minor 28
fringillinus, Parus 16
fringilloides, Lonchura 6
frontalis, Nonnula 118
Fulica cristata 12
fuligula, Ptyonoprogne 17-19
Fulmarus glacialoides 34, 58-59
furcata, Thalurania 118
fuscata, Sterna 29, 100
fuscescens, Dendropicos 16
fuscicollis, Calidris 58, 63
galactotes, Cisticola 12-13
Galbula ruficauda 118
Galerida modesta 144
— theklae 139-142
Galerida theklae harrarensis, subsp. nov.
139
Galerida theklae huei, subsp. nov. 141
galeritus, Anorrhinus 172
Gallinago gallinago 174
— media 174
— nigripennis 68, 174
gallinago, Gallinago 174
Gallinula chloropus 12
Gallus gallus 28
gallus, Gallus 28
gambensis, Plectropterus 12
Garrulax erythrocephalus 172
— mitratus 171
garrulus, Coracias 126-130
Geopelia cuneata 105
georgica, Anas 58, 61
Geospiza fortis 77
Geothlypis trichas 77
Geranoaetus melanoleucus 62
Geronticus calvus 163
— eremita 163
giganteus, Macronectes 34, 58
glacialoides, Fulmarus 34, 58-59
Glaucidium brodiei 172
— tephronotum 25-26
Glaucis hirsuta 118
glaucurus, Eurystomus 16-17, I 3°
goertae, Mesopicos 16
goisagi, Gorsachius 176
Gorsachius goisagi 176
— melanolophus 176
Gracula religiosa 171
grandis, Bradypterus 169
graueri, Bradypterus 167-169
gravis, Puffinus 58-59
griseicauda, Treron 119
griseopyga, Pseudhirundo 138
griseus, Puffinus 58-59
guildingii, Amazona 120-123
gularis, Monticola 149
Gygis alba 30
gymnocephalus, Picathartes 23-24
Gypaetus barbatus 65, 68, 161-162
Haematopus ater 58, 62
— leucopodus 58, 63
— palliatus 58, 62
hagedash, Hagedashia 12
Hagedashia hagedash 12
Halcyon albiventris 15
— chelicuti 16
— chloris 80, 158
— coromanda 80
— pileata 80
— sancta 79-80
— senegalensis 79
Harpactes duvauceli 172
harrarensis, Galerida 139
hartlaubi, Tauraco 15
Hartlaubius auratus 153
Herpetotheres cachinnans 1 1 7
Hesperornjs 123-125
hildebrandti, Francolinus 14
Himantopus himantopus 40
himantopus, Himantopus 40
Hippolais polyglotta 102
hirsuta, Glaucis 118
hirundinacea, Sterna 36, 58-59, 62-63
Hirundo 20
— abyssinica 17
— senegalensis 17-18
— smithii 17-18
holomelaena, Psalidoprocne 15, 69
Hoplopterus armatus 14-15
hordeaceus, Euplectes 161
hortulana, Emberiza 81
horus, Apus 17-18
huei, Galerida 141
hunteri, Cisticola 64, 66, 70
Hydranassa tricolor 1 1 7
Hylobates syndactylus 1 71-172
Hyphanturgus ocularis 14-15
hypoleucos, Pomatorhinus 172
— Tringa 39-40
Ichthyornis 123-126
— dispar 124-125
imerina, Cossypha 145
— Monticola 145, 152
importunus, Andropadus 4
incana, Tringa 29
inda, Chloroceryle 118
insularis, Sarothrura 130-133
intermedius, Ploceus 14
islandica, Bucephala 6-9
Ispidina picta 82-83
Jacamerops aurea 118
Jacana jacana 117
jacana, Jacana 117
jamaicensis, Leptotila 106
— Oxyura 98-99
jardineii, Turdoides 24
javensis, Dryocopus 171
johnstoni, Nectarinia 66-67, 7°
kenricki, Stilbopsar 15
kilimensis, Nectarinia 17-19
lacrymosum, Tricholaema 16
lacteus, Bubo 13, 26
Lagonosticta rubricata 14
Lalage nigra 1 5 8
Laniarius aethiopicus 16-19
Lanius collaris 16-19, 7°
— collurio 92-96
lapponica, Limosa 100
Larus dominicanus 33, 36, 58-59, 63
— maculipennis 58-59, 63
larvatus, Oriolus 16
Laterallus albigularis 117
laticauda, Coliuspasser 14
lcadbeateri, Bucorvus 89-91
Leptotila cassinii 117
— jamaicensis 106
— rufaxilla 106
— verreauxi 1 17
lcpturus, Phaethon 28
Lessonia rufa 36
leucocephala, Columba 107
leucocephalus, Lybius 16
leucogaster, Cinnyricinclus 1 60-1 61
— Sula 28
leuconotus, Thalassornis 12-13
Leucophaeus scoresbii 35, 58, 63
leucopodus, Haematopus 58, 62
leucoptera, Pterodroma 27
leucopterus, Platysmurus 1 71-172
leucopygius, Turdoides 24
leucorodia, Platalea 124
leucotis, Buccanodon 15
Limnocorax flavirostra 12
Limosa lapponica 100
lineatum, Tigrisoma 117
livia, Columba 38-39, 107
Locustella fluviatilis 4, 6
— naevia 101
Lonchura fringilloides 6
longicauda, Psittacula 171
longirostris, Rhizothera 172
longuemareus, Phaethornis 118
Lophonetta specularoides 58, 61
loricatus, Celeus 118
ludovicianus, Pheucticus 77
lugens, Sarothrura 132-133
— Streptopelia 17
lunata, Sterna 29
luteicapilla, Euphonia 77
Lybius leucocephalus 16
— melanopterus 15
maccoa, Oxyura 98-99
macrodactyla, Napothera 171
Macronectes giganteus 34, 58
Macronyx 18
— aurantiigula 14-15, 19, 70
— croceus 144
Macropygia ruficeps 172
macrorhynchos, Notharcus 118
macrurus, Caprimulgus 172
macularia, Actitis 117
maculata, Stachyris 171
maculipennis, Larus 58-59, 63
maculosa, Columba 107
— Prinia 153
madaga scarier) sis, Rallus 133
magellanicus, Phalacrocorax 34, 58, 60
— Spheniscus 33-34, 58-59
magnirostris, Buteo 117
major, Crotophaga 118
— Podiceps 58-59
malabaricus, Copsychus 1 71-172
malacensis, Anthreptes 159
— Polyplectron 172
Malaconotus zeylonus 1 5 3
Malacopteron affine 171
Malacoptila panamensis 118
malayanus, Anthracoceros 171
martii, Baryphthengus 118
martinica, Porphryula 36-37
mauri, Ereunetes 117
maxima, Collocalia 171
maximus, Saltator 77
— Thalasseus 58, 63
mbangensis, Cisticola 144
media, Gallinago 174
mediocris, Cinnyris 15
Megalaima australis 171
— oorti 171
Megaloprepia 120
melanocoryphus, Cygnus 35
melanoleucos, Campephilus 118
melanoleucus, Geranoaetus 62
melanolophus, Gorsachius 175
melanophris, Diomedia 34, 58-59
melanopterus, Lybius 15
melanotis, Coccopygia 14-15
melanotos, Calidris 3, 6
melanurus, Trogon 1 1 8
melba, Apus 65, 71
Melittophagus bullockoides 16
— oreobates 15
— pusillus 16
Melocichla mentalis 14, 19
mentalis, Melocichla 14, 19
mercierii, Ptilinopus 31
Mesopicos goertae 16
micropterus, Cuculus 171
migrans, Milvus 70
milanjensis, Arizelocichla 15
Milvago chimachima 117
Milvus migrans 70
miniatus, Myioborus 77
minor, Fregata 28
minuta, Columbina 117
— Piaya 118
minutus, Numenius 99-100
Mirafra africana 14-15
miriflcus, Rallus 115
mitrata, Numida 14
mitratus, Garrulax 171
modesta, Galerida 144
modestus, Zonibyx 58
molitor, Batis 16-19
momota, Momotus 1 1 8
Momotus momota 118
Monasa morphoeus 118
Monticola angolensis 154
— bensoni 145
— cinclorhynchus 149
— gularis 149
— imerina 145, 152
— rupestris 154
— saxatilis 81
— sharpei 145-154
— solitarius 81
morio, Onychognathus 16
morphoeus, Monasa 118
Motacilla aguimp 16
— alba 97-98
— clara 12-13
mozambicus, Serinus 144
Muscicapa 101
mutata, Tchitrea 153
Myiarchus ferox 114
Myioborus miniatus 77
Myiodynastes 115
myoptilus, Apus 70
Myrmecocichla nigra 144
naevia, Coracias 127-130
— Locustella 101
nagtglasii, Buceros 120
namaquus, Thripias 16-17
nana, Turnix 139
Napothera macrodactyla 171
natalensis, Chloropeta 14
— Cisticola 144
— Cossypha 43
nativitatis, Puffinus 28
nebularia, Tringa 68
Nectarinia cuprea 144
— famosa 153
— johnstoni 66-67, 7°
— kilimensis 17-19
— tacazze 15
neglecta, Pterodroma 27
Nesillas typica 153
Nesofregetta albigularis 28
newtoni, Calamocichla 133
— Falco 108-109
niansae, Apus 17-18
nicobarica, Caloenas 53
niger, Chlidonias 117
nigra, Lalage 158
— Myrmecocichla 144
nigriceps, Arizelocichla 15
— Spermestes 14
nigricollis, Busarellus 117
— Stachyris 171
nigripennis, Gallinago 68, 174
nigrirostris, Columba 117
Nigrita canicapilla 15
nigrocincta, Tangara 78
Nilaus afer 16
Nonnula frontalis 1 1 8
— ruficapilla 118
Notharctus macrorhynchos 118
— tectus 1 1 8
nubica, Campethera 16, 72-74
Numenius minutus 99-100
Numida mitrata 14
Nyctanassa violacea 117
Nycticorax caledonicus 175
— nycticorax 58, 61, 175-176
nycticorax, Nycticorax 58, 61, 175-176
Nyctidromus albicollis 118
Nyctiornis amicta 171
Nystalus r adiatus 1 1 8
occipitalis, Podiceps 58-59
oceanicus, Oceanites 58-59
Oceanites oceanicus 58-59
ochrocephala, Amazona 118
ocularis, Hyphanturgus 14-15
olivacea, Cyanomitra 15-16
— Piranga 77, 156
olivaceus, Phalacrocorax 58-59
— Picumnus 118
— Turdus 15
Onychognathus morio 16
— tenuirostris 70
— walleri 15
oorti, Megalaima 171
oreobates, Melittophagus 15
oreophilus, Buteo 13, 65, 68
Oriolus larvatus 16
— xanthonotus 172
Ortalis cinereiceps 117
Orthotomus cucullatus 172
— sepium 171
Othyphantes reichenowi 14-15
Oxyura australis 98
— jamaicensis 98-99
— maccoa 98-99
Pachy coccyx audeberti 1 60-1 61
Pachyptila 58
— belcheri 59
pacifica, Ducula 53
pacificus, Puffinus 27
palliatus, Haematopus 58, 62
palmarum, Thraupis 75, 156
paludicola, Riparia 17-18
panamensis, Malacoptila 118
papa, Sarcoramphus 117
papuensis, Coracina 159
paradiseus, Dicrurus 171
paradisi, Terpsiphone 172
Parus afer 1 5 3
— fringillinus 16
Passerina cyanea 77
patachonicus, Tachyeres 35, 58, 61
pectoralis, Rallus 115
pecuarius, Charadrius 14
Pelargopsis capensis 158
penicillata, Eucometis 77
pensylvanica, Dendroica 77
peregrinus, Falco 13, 58, 61, 108
Pernis apivorus 116
— celebensis 116
perspicillata, Pulsatrix 118
petechia, Dendroica 77
petiti, Campephaga 41
Petrochelidon albifrons 20
— pyrrhonota 20
Petrocincla 145
Peucedramus 74
phaeocephalus, Criniger 171
Phaethon aethereus 28
— lepturus 28
— rubricauda 28
Phaethornis longuemareus 118
— superciliosus 118
Phalacrocorax albiventer 34-35, 58, 60
— atriceps 35, 58, 61
— bougainvillii 34-35
— magellanicus 34, 58, 60
— olivaceus 58-59
Phasianus 53
Pheucticus ludovicianus 77
phoenicea, Campephaga 41, 44-47
Phoenicopterus chilensis 35, 58, 61
Phoeniculus purpureus 16
Pholia sharpii 15
Phyllastrephus strepitans 4
Phyllolais pulchella 16
Phylloscopus sibilatrix 5-6
— trochilus 5, 102
Piaya cayana 118
— minuta 118
Picathartes gymnocephalus 23-24
picazuro, Columba 105, 107
picta, Ceyx 82, 84-85
— ■ Chloephaga 58, 61
— ■ Ispidina 82-83
Picumnus olivaceus 118
pileata, Halcyon 80
Piranga erythromelas 76
— olivacea 77, 156
Pitta erythrogaster 32-33
Platalea leucorodia 124
platypterus, Buteo 117
platyrhynchum, Electron 118
Platysmurus leucopterus 171
paludicola, Riparia 17-18
Plectropterus gambensis 12
Ploceus castaneiceps 14-15
— intermedius 14
— spekei 14
plumbea, Columba 103-107
Pluvialis dominica 28
Pluvianellus socialis 58, 63
Podiceps dominicus 117
— major 58-59
— occipitalis 58-59
— rolland 58-59
podiceps, Podilymbus 117
Podilymbus podiceps 117
Poliocephalus ruficollis 12-13
poliocephalus, Caprimulgus 14, 19, 71
polyglotta, Hippolais 102
Polyplectron malacensis 172
Pomatorhinus hypoleucos 172
Pomatostomus superciliosus 24
— temporalis 24
pompadora, Treron 119
Porphyrula martinica 36-37
pretiosa, Claravis 117
Prinia maculosa 153
— subflava 14, 18-19, J 44
Prionops retzii 161
Procellaria aequinoctialis 58-59
Procelsterna cerulea 29
Procnias 75
Prosobonia cancellatus 28
Psalidoprocne holomelaena 15, 69
Pseudhirundo 20
— griseopyga 138
psilolaemus, Francolinus 70
Psilopogon pyrolophus 172
Psittacula longicauda 171
Pterodroma alba 27
— arminjoniana 27
— leucoptera 27
— neglecta 27
— rostrata 27
Pter oglossus torquatus 1 1 8
Ptilinopus bernsteinii 120
— coralensis 30-31
— dupetithouarsii 31
— formosus 120
— mercierii 31
— purpuratus 31
— superbus 31
Ptyonoprogne fuligula 17-19
pucherani, Centurus 118
Puffinus gravis 58-59
— griseus 58-59
— nativitatis 28
— pacificus 27
pulchella, Phyllolais 16
pulchra, Sarothrura 135
pulchrirostris, Buceros 119-120
Pulsatrix perspicillata 118
punctata, Anas 12
punctuligera, Campethera 72-74
purpuratus, Ptilinopus 31
purpureus, Phoeniculus 16
pusillus, Melittophagus 16
Pycnonotus barbatus 4, 16-19
— zeylanicus 172
pyrolophus, Psilopogon 172
pyrrhonota, Petrochelidon 20
Quelea 15, 69
quiscalina, Campephaga 41
radiatus, Nystalus 118
Rallus madagascariensis 133
— mirificus 1 1 5
— pectoralis 115
— striatus 115
rapax, Aquila 13
Recurvirostra avosetta 40
regulorum, Balearica 12-13
reichenowi, Othyphantes 14-15
religiosa, Gracula 171
reaauldi, Carpococcyx 174
retzii, Prionops 161
rhinoceros, Buceros 172
Rhinopomastus cyanomelas 16
Rhizothera longirostris 172
Riparia 138
— paludicola 17-18
rolland, Podiceps 58-59
Rollulus rouloul 53-54
rostrata, Pterodroma 27
Rostrhamus sociabilis 117
rouloul, Rollulus 53-54
rubidiceps, Chloephaga 61
rubricapillus, Centurus 118
rubricata, Lagonosticta 14
rubricauda, Phaethon 28
ruckeri, Threnetes 118
rufa, Lessonia 36
— Sarothrura 131, 135
rufaxilla, Leptotila 106
ruficapilla, Nonnula 118
ruficauda, Galbula 118
ruficeps, Macropygia 172
ruficollis, Poliocephalus 12-13
rufipennis, Butastur 129
rufofuscus, Buteo 13, 19, 65
rufus, Trogon 1 1 8
rumicivorus, Thinocorus 35, 58, 63
rupestris, Monticola 154
sacra, Egretta 28
Saltator maximus 77
sancta, Halcyon 79-80
Sarcoramphus papa 117
Sarothrura affinis 70, 133
— ayresii 132
— boehmi 131
— elegans 134-135
— insularis 130-133
— lugens 132-133
— pulchra 135
— rufa 131, 135
— watersi 132
saularis, Copsychus 172
Sauropatis 79-80
saxatilis, Monticola 81
Saxicola torquata 14-15, 18-19, 65, 69
sayaca, Thraupis 77
scirpaceus, Acrocephalus 102-103
Scopus umbretta 109
scoresbii, Leucophaeus 35, 58, 63
scriptoricauda, Campethera 72-74
Seicercus umbrovirens 1 5
Selenidera spectabilis 1 1 8
semipalmatus, Catoptrophorus 117
semirufa, Cossypha 15
semitorquata, Streptopelia 17, 19
senegala, Tchagra 16, 144
senegalensis, Alcedo 79
— Centropus 173
— Chalcomitra 17-18
— Ephippiorhynchus 12-13
— Halcyon 79
— Hirundo 17-18
— Stigmatopelia 17
sepiarium, Trichastoma 24
sepium, Orthotomus 171
Serinus burtoni 15
— canicollis 67
— flavivertex 17-18
— mozambicus 144
— striolatus 17-19, 64, 67-68, 70
serriana, Coua 3
sharpei, Cossypha 147
— Monticola 145-154
sharpii, Bycanistes 55
— Pholia 15
shelleyi, Francolinus 70
sibilatrix, Phylloscopus 5-6
similis, Anthus 14, 18-19, 8l
siparaja, Aethopyga 159
skua, Catharacta 35, 58, 63
smithii, Hirundo 17-18
sociabilis, Rostrhamus 117
socialis, Pluvianellus 58, 63
solitaria, Tringa 117
solitarius, Monticola 81
sordida, Cercomela 64-66, 70
sparsa, Anas 12, 70
sparveriodes, Cuculus 171
spectabilis, Selenidera 118
specularoides, Lophonetta 58, 61
spekei, Ploceus 14
Spermestes cucullatus 14
— nigriceps 14
Spheniscus demersus 34
— magellanicus 33-34, 58-59
Spilornis cheela 171
Spizaetus cirrhatus 172
Spreo bicolor 152
squamatus, Francolinus 14
Squatarola squatarola 100
squatarola, Squatarola 100
Stachyris chrysaea 172
— maculata 171
— nigricollis 171
Stephanibyx 68
— coronatus 14
Sterna bergii 29
— dougalii 21
— eurygnatha 36
— fuscata 29, 100
— hirundinacea 36, 58-59, 62-63
— lunata 29
— trudeam 58, 63
Stigmatopelia senegalensis 17
Stilbopsar kenricki 15
stolidus, Anous 29-30
strepitans, Phyllastrephus 4
Streptopelia 105
— capicola 17
— lugens 17
— semitorquata 17, 19
Streptoprocne 196
striatus, Colius 16, 18-19
— Rallus 115
striolatus, Serinus 17-19, 64, 67-68, 70
subbuteo, Falco 99-101, 127
subflava, Prinia 14, 18-19, x 44
subtilis, Buteogallus 117
subvinacea, Columba 104
Sula leucogaster 28
— sula 28
sula, Sula 28
sulcirostris, Crotophaga 118
sulfureopectus, Chlorophoneus 16, 18-19
sulphurata, Campephaga 16
superbus, Ptilinopus 31
superciliosus, Centropus 14
— Phaethornis 118
— Pomatostomus 24
Sylvia borin 102-103
— cantillans 101
Sylvietta whytii 16, 19
syndactylus, Hylobates 1 71-172
tacazze, Nectarinia 15
Tachyeres patachonicus 35, 58, 61
tahapisi, Emberiza 81
taitensis, Eudynamis 52
talpacoti, Columbina 1 1 7
Tanagra violacea 156-157
Tangara 76-77, 155
— nigrocincta 78
Tauraco hartlaubi 15
Tchagra australis 16, 18-19
— senegala 16, 144
Tchitrea mutata 153
— viridis 15-16, 18-19
tectus, Notharctus 118
temmin.ckii, Eurostopodus 171
temporalis, Pomatostomus 24
tenuirostris, Anous 29-30
— Onychognathus 70
tephronotum, Glaucidium 25-26
Terathopius ecaudatus 13
Terpsiphone paradisi 172
Tersina viridis 75-78, 155-157
Thalasseus maximus 58, 63
Thalassornis leuconotus 12-13
Thalurania furcata 118
theklae, Galerida 139-142
Thinocorus 75
— rumicivorus 35, 58, 63
Thraupis 1 5 5
— episcopus 77
— palmarum 75, 156
— sayaca 77
Threnetes ruckeri 118
Thripias namaquus 16-17
Tigrisoma lineatum 117
tiphia, Aegithina 158-159
Tockus 171
— alboterminatus 15, 55
— flavirostris 55
torquata, Ceryle 118
— Saxicola 14-15, 18-19, 65, 69
tor quatus, Pteroglossus 1 1 8
Treron aromatica 119
— australis 17
— curvirostra 158
— griseicauda 119
— pompadora 1 1 9
trichas, Geothlypis 77
Trichastoma abbotti 172
— bicolor 24
— sepiarium 24
Tricholaema lacrymosum 16
tricollaris, Charadrius 14
tricolor, Hydranassa 1 1 7
trigonostigma, Dicaeum 159
Tringa flavipes 117
— hypoleucos 39-40
— incana 29
— nebularia 68
— solitaria 117
trivialis, Anthus 81
trochilus, Phylloscopus 5, 102
Trogon melanurus 118
— rufus 118
— viridis 118
trudeaui, Sterna 58, 63
Turdoides jardineii 24
— leucopygius 24
Turdus olivaceus 1 5
Turnix 75
— nana 139
Tympanistria tympanistria 15
tympanistria, Tympanistria 15
typica, Nesillas 153
Tyto alba 26
umbretta, Scopus 109
umbrovirens, Seicercus 1 5
Upucerthia dumetaria 36
urubitinga, Buteogallus 117
Vanellus chilensis 35, 58
vasa, Coracopsis 153
venustus, Cinnyris 17-19
verreauxi, Leptotila 1 1 7
— Aquila 70, 166
violacea, Nyctanassa 117
— Tanagra 156-157
virescens, Butorides 117
viridis, Tchitrea 15-16, 18-19
— Tersina 75-78, 155-157
— Trogon 118
viridissima, Aegithina 158
vocifer, Cuncuma 13
walleri, Onychognathus 15
watersi, Sarothrura 132
whytii, Sylvietta 16, 19
wilsonia, Charadrius 117
woodfordii, Ciccaba 26
woosnami, Cisticola 14, 19
xanthonotus, Oriolus 172
Xanthopsar flavus 172
yokanae, Bradypterus 167
zeylanicus, Pycnonotus 172
zeylonus, Malaconotus 153
Zonibyx modestus 58
zoniventris, Falco 1 08-1 11
Zonotrichia capensis 77
Zosterops eurycricotus 15
Corrigenda
p. ii, line 47: 'rising', not 'rising'
p. 13, line 25: i Motacilla\ not 'Moticilld
p. 14, table 4: l hildebrandti'' not 'hz/debrantf
p. 15, table 5 : i Stilbopsar\ not 'Stilbospar' ; 'medzocris' not 'mediocria .
p. 15, last line: 'observation', not 'obesrvation' ; 'carried', not 'varried'.
p. 16, table 6: i lacrymosum\ not i lachrymosum''
p. 17, table 7: i Ptyonoprogne\ not ' ' Pytonoprogne
p. 19, line 30: l Carduelis\ not ' Cardeuelis*
p. 23, line 22: 'nesting', not 'nestling'
p. 29, line 13: 'bergz'f, not 'bergf
p. 34, line 49: i boHgamvillii\ not 'bougamvzl/zz'
pp. 36-37: 'Olson', not 'Ohlson' (in paper by Frost & Siegfried)
p 43, line 33 : 'east of, not '33 to'
p. 43, line 37: 'Amani', not 'Amari'
p. 44, line 20: '356', not '351'
p. 45, line 24: 'May', not 'Mat'
p. 53 : In title to paper by Farrand & Olson, 'Malaysian', not 'Maylaysian'
p. 65, line 21: 'Rodents', not 'Rodent'
line 9 : To re-read 'same time, and integration is possible. Attention is drawn to'
p. 97 : In title to paper by Larmuth, 'alba, not l alhd
p. 153, line 27: i Tchitrea\ not 'Tchitrd
p. 172, line 8: l aeneus\ not 'aenus'
The Caxton & Holmesdale Press, Sevenoaks
r *
Bulletin of the
British Ornithologists' Club
Edited by
C. W. BENSON
Volume 93 No. 1
March 1973
Committee
Sir Hugh Elliott, Bt., o.b.e. {Chairman)
J. H. Elgood {y ice-Chair man)
C. W. Benson, o.b.e. {Editor)
R. E. F. Peal {Hon. Secretary)
P. Tate {Hon. Treasurer)
J. K. Adams
Dr. P. J. K. Burton
P. Hogg
C. J. Mead
Bulletin of the
BRITISH ORNITHOLOGISTS' CLUB
Vol. 93 No. 1 Published: 20th March, 1973
The six hundred and eightieth meeting of the Club was held at the Cafe
Royal, 69 Regent Street, London, W.i, on Tuesday, 16th January, 1973 at
7 p.m.
Chairman: Sir Hugh Elliott, Bt., o.b.e.; present 14 members and 8 guests.
The speaker was Mrs. M. K. Rowan, who gave an illustrated address on
African brood parasites.
The Annual General Meeting of the Club will be held at the Cafe Royal,
68 Regent Street, London, W.i on Tuesday, 8th May, 1973 at 6 p.m.
AGENDA
1. Minutes of the last Annual General Meeting.
2. Report of the Committee and Accounts for 1972.
3. The Bulletin. L,
4. Election of Officers :
(a) The Committee proposes that Mr. R. F. Peal be re-elec
Secretary.
(b) The Committee proposes that Mr. P. Tate be re-elected Honorary
Treasurer.
(c) The Committee proposes that Mr. J. H. R. Bos wall be elected a
member of the Committee vice Mr. J. K. Adams.
by order of the Committee,
RONALD E. F. PEAL,
Honorary Secretary.
Report of the Committee for 1972
In January, March and May, the meetings of the Club were held at the
Criterion, in Piccadilly Circus, but with the closing-down of the Criterion
were transferred, for the July, September and November meetings, to the
not far distant Cafe Royal. The total number of members and guests
attending showed a welcome increase to 191 (138 in 197 1), the highest since
1967 (193). This may be a reflection of the convenient situation and good
value of the present venue; it is hoped that attendance will continue at the
present level or show a further increase in 1973, since the favourable dinner-
charge of £1.75 per head including service is partly dependent on that
factor.
The Committee regrets to report the death of one member during the
year, Mr. R. E. Heath. Ten new members joined the Club, but there were
six resignations and the membership of 17 members has had to be terminated
under Rule 4, a nett loss of 14 members. This may be regarded as a further,
but it is to be hoped final, reaction to the increased subscription introduced
in 1 97 1, the total loss of membership over the two years being about 10%.
It is important that this should now be made good as quickly as possible,
since a membership of the order of 275 (about 50 more members than at
present), combined if possible with some increase in the number (at present
about 80) of non-member subscribers, would certainly be the surest means of
achieving financial stability.
At the Annual General Meeting on 9th May, Mr. P. Hogg was elected to
the Committee in place of Mrs. J. D. Bradley, who retired by rotation. A
major change approved by the Meeting and implemented by a decision of the
Committee in October was that publication of the Bulletin (which in 1972
still comprised 172 pages and, technically, six issues, although there was in
fact a 'June/ August' double number) should be placed on a quarterly basis
in 1973. One result of this economy measure is that the Audited Accounts for
1972 cannot be circulated with the present Report, but will have to be tabled
separately at the 1973 Annual General Meeting. It is proposed, however, to
revert in future years to the pre- 1969 practice of publishing the Report and
Accounts, as well as the Minutes of the A.G.M., in the Bulletin. The four
pages used for this purpose will be additional, thus increasing the length of
each annual volume of the Bulletin to 176 pages.
In general the Club's financial position showed a further slight improve-
ment in 1972, but this, as explained at the Annual General Meeting (see
Bulletin Vol. 92: 3 & 4), was largely due to the completion of the sorting of
Bulletin stocks and the consequent resumption of back number sales (which
included a considerable backlog of orders). It would be unrealistic to suppose
that this level of sales can be maintained in future years, so that as already
indicated some increase of membership, supplemented by the continuing
exercise of the strictest economy (as exemplified by the quarterly publication
of the Bulletin), remain essential.
A specimen of Coua delalandei (Temminck) (Cuculidae)
in the Naturhistorisches Museum, Vienna (Austria)
by Herbert Schijter
Received 26th September 1972
Benson & Schiiz (1971) have mentioned the existence of a specimen of Coua
delalandei in the Staatliches Museum fur Naturkunde, Stuttgart, not listed by
Greenway (1967). There is also a mounted specimen of this species, believed j
to be extinct, in the Naturhistorisches Museum, Vienna. Neither was this
specimen listed by Greenway (1967), nor by Luther (1970), although it was
included by Sassi (1939) in his contribution on rare birds in the Vienna
Museum.
The Vienna specimen oiCoua delalandei, no. NMW 50810, was bought by
Johann Natterer in 1840 during his journey to France and England, when he
visited museum collections and dealers in natural history objects (Roki-
tansky 1957). Its acquisition number is 1844.I.252 in the collection brought
back by Johann Natterer, comprising more than 500 specimens acquired
from 28 different sources. At this time (1844) Joseph Natterer, the elder
brother of Johann, was curator of the ornithological collections in the
museum. Unfortunately the specimen does not bear an exact date of collec-
ting. It is merely marked "1840", with the epithets "Evans" and "Madagas-
car". The name Evans does not appear in the list of specimens acquired,
although it is mentioned at the end of the list together with the names of
other dealers, with the remark that seven specimens were acquired from this
source. So it is not possible to ascertain which are the other six specimens
obtained from Evans, although it is likely that other specimens from
Madagascar in the collection were obtained from other dealers, such as
Verreaux. The species are listed in a systematic sequence, so that the
specimens from various sources are mixed. The date 1840 on the label of the
C. delalandei cannot be regarded as that of collecting, but merely that of
acquisition. Nevertheless it is possible that this specimen, and the one in
Stuttgart, were collected later than 1834, although neither Jouanin (1962)
nor Greenway (1968) could find evidence of one obtained later. The Vienna
specimen is still in good condition, even though in the past it has undoubtedly
been on public exhibit.
In the list of acquisitions another specimen appears under the name of this
species, no. 1844.I.25 3. Sassi (1939) referred to it as having been perhaps lost
or given away in exchange. I have ascertained that it is still in the collection.
It is not however a Coua delalandei but a C. serriana. It was not identified to
species originally, the name C. serriana having been added on the label later.
As Pucheran's description of C. serriana is dated 1845, the specimen was
received in Vienna before the species had been named. The name "Richard"
appears on the original label, so that it does not appear to have come from
the same source as the C. delalandei. Although it was without doubt originally
mounted, it is now kept as a study skin.
To summarise, attention is re-drawn to the existence of a specimen of
Coua delalandei in the Naturhistorisches Museum, Vienna, but there is no
evidence that a second specimen was ever acquired, as has been claimed.
References :
Benson, C. W. & Schuz, E. 1971. A specimen oi Coua delalandei (Temminck) (Cuculidae).
Bull. Brit. Orn. CI. 91: 159-160.
Greenway, J. C. 1967. Extinct and vanishing birds of the world. Revised ed'n. New York:
Dover Publications.
Jouanin, C. 1962. Inventaire des oiseaux eteints ou en voie d'extinction conserves au
Museum de Paris. Terre Vie 109: 257-301.
Luther, D. 1970. Die ausgestorbenen vogel der welt. Wittenberg Lutherstadt: Neue Brehm
Biicherei no. 424.
Rokitansky, G. 1957. Johann Natterer, Erster Ornithologe Oesterreichs. Jonrn. Ornith. 98:
133-144.
Sassi, M. 1939. Die wertvollsten Stucke der Wiener Vogelsammlung. Ann. Naturhistor.
Mus. Wien 50: 395-409.
Six species of birds new to Ethiopia
by J. S. Ash
Received 28th September 197$
Three of the following six species, not previously recorded in Ethiopia, have
been discovered since the publication of the recent Checklist (Urban &
Brown 1971); the other three were included just prior to going to press.
Calidris melanotos^ American Pectoral Sandpiper: One was observed
feeding at dawn along the western shore of the artificial Rift Valley lake,
formed by a dam, at Koka (8° 27' N, 39 6' E), Shoa Province, Ethiopia, on
1 8th March 1971. It had not been seen on the previous 13 days. I had another
opportunity to look for it in the late afternoon, when it was found again in
the same place and collected. It proved to be a male with regressed gonads,
with wing-length 148 mm and weight 67.4 g. The specimen is deposited
at the Smithsonian Institution, where Dr. George E. Watson has kindly
confirmed the identification. There are now ten examples of this Nearctic
species to be recorded on the African continent, and eight of them are from
south of the Sahara. The previous records have been one each in Kenya,
Botswana and Rhodesia, two in Morocco and four in South Africa; all since
1949 (Ginn & Brooke 1971 ; Kemp 1972).
It seems likely that this and the other birds from south of the Sahara were
displaced across the north Atlantic on autumn passage and continued south
to overwinter in tropical Africa. If such displaced birds return north in the
Palearctic spring, they could account for some of the spring occurrences in
Europe. Excluding the two Moroccan records, it is of interest that the others,
except for one bird in South Africa in December, occurred in March-May.
Chlorocichla flavicollis, Yellow-Throated Leaf-Love: The Pycnonotidae, of
which about 3 3 species are currently recognized in East Africa, is a poorly
represented family in Ethiopia. Until the present species was found, only
three others were known: Pycnonotus barbatus common and widespread,
A.ndropadus importunus in the south-east and Phyllastrephus strepitans common
in parts of the south (Urban & Brown 1971). It is possible that others remain
to be identified in the south-western forests (Brown & Urban 1970) and
along the Wabi Shabelli in southern Ogaden (personal observations).
The present species was first found at Didessa (9 02' N, 3 6° 09' E),
Wollega Province, in a mist net on 25 th July 1971. Since then six further
examples have been caught as follows : —
Date
Wing {mm)
Weight
'W
Time
Wing Moult
25 July 70
106
44.9
1600
Nil
18 February 71
103
38.6
2130
23 February 71
104
45.2
1 100
6 July 71
107
45.8
1130
7 July 7 1
104
37-7
0700
8 July 71
94
38.8
1700
Nil
9 July 71
98
40.7
1400
6 February 72*
94
37-0
0830
Nil
*Retrap of the bird caught on 8th July 1971.
All the birds were caught along the sides of two small streams, along which
there are a few trees and a thick underscrub, on the edge of the plateau above
the Didessa River gorge. The species could be quite widespread in the
abundant similar habitat in this region.
Two specimens examined at the British Museum by Mrs. B. P. Hall and
Mr. Derek Goodwin are identified as C. f. soror but are stated to be greener
on the back than neighbouring forms and show some approach to palli-
digula y being a very little more yellowish on throat and underparts than most
of the specimens of soror in the British Museum.
The Didessa birds represent a considerable extension of the known range
of the species, north-eastwards into the Ethiopian highlands from Uganda.
Locus tella fluviatilis, River Warbler: Urban & Brown (1971) stated that the
River Warbler "should occur in Ethiopia but has not been recorded as yet".
It is therefore of interest to record five examples in two autumns at Lake
Abiata (7 36' N, 3 8° 40' E) in the Ethiopian Rift Valley. Three of these were
collected and one each deposited in the collections of the British Museum
and Smithsonian Institution. The records are as follows: —
Date
Age
Wing {mm) Weight {g)
Time
Wing Moult
29 October 70
1 Y
73
17.6
1730
Nil
30 October 70
p
77
18.5
1030
Nil
31 October 71
1 Y
69
14.2
1 100
Nil
13 October 71
?
69
14.8
1 100
Nil
22 October 71
?
74
18.0
1030
Nil
The timing of migration is of interest. In 1970 the area was worked from
10th October to 2nd November, and the birds were captured in the last
three days of October. The 1971 records, when the area was worked from
1 2th October to 2nd November, were much earlier. R. J. Dowsett informs
me (in litt.) that arrivals in Kenya seem to be about mid-December, and in
Zambia later in the month, so that there may be an interval of about two
months before these birds traverse the relatively short distance to their
wintering areas. So far there is no evidence that this species occurs in
Ethiopia in mid winter, in spite of extensive work in suitable habitat. The
Abiata birds were found in thick herbaceous growth adjoining a river and
lake.
Phylloscopus slbllatrlx, Wood Warbler : It is curious that the Wood Warbler
has not been recorded previously in Ethiopia (Urban & Brown 1971) for
the species is well-known further west and south (Mackworth-Praed &
Grant 1955) and at a roughly similar latitude in West Africa (e.g. Dowsett &
Fry 1971). A specimen was netted, ringed and released, on 1st May 1971 in a
small area of deciduous woodland located on the periphery of a flood plain
of the River Awash where it flows through the Danakil Desert. The area is
known as Bahadu (io° 05' N, 40 37' E), near Gewani (= Gauani), Harar
Province in eastern Ethiopia.
It was easily identified among the few Willow Warblers Phylloscopus
trochilus by the moss-green coloration of its upper parts and yellow breast
demarcated from the white of the rest of the underparts. Its diagnostic wing
formula (short 1st primary, 2nd=4th, 3rd longest, 3rd and 4th only emar-
ginated) confirmed this, and it weighed 11. 2 gat 1130; wing-length 71 mm.
Acrocephalus baetlcatus, African Reed Warbler: The African Reed Warbler
was included in the recent Checklist (Urban & Brown 1971) on the basis of a
specimen I collected in 1970. There are now records of six more birds from
two localities, and it seems appropriate to summarize these here as they
represent a notable eastwards extension of the species' known range.
Summary of records in chronological order:
Date
Wing (mm)
Weigh
Hi)
Time
Locality
zz April 70
55
7-4
0930
Bahadu
13 March 71
55
7-3
1630
Koka
3 May 71
55
8.0
1630
Bahadu
6 May 71
56
8.1
0800
Bahadu
7 May 71*
55
8.1
0800
Bahadu
23 November 71
57
8.0
1030
Bahadu
27 March 72
56
7-9
0730
Koka
2 April 72
58
7-9
0830
Koka
4 April 72
56**
7-5
0730
Koka
*Retrap of the bird caught on 3rd May 1971.
**Retrap of the bird caught on 27th March 1972.
Bahadu (io° 05' N, 40 37' E) is an area of permanent marshland on a
flood-plain on the River Awash, where it flows through the western Danakil
desert in eastern Ethiopia, close to Gewani in Harar Province. The area at
Koka (8° 27' N, 39 06' E), in the Rift Valley in Shoa Province, is thick
lakeside acacia bush.
The first Ethiopian specimen was deposited in the British Museum where
it was determined by Mrs. B. P. Hall as agreeing with the western form A. b.
clnnamomeus. Prior to the discovery of the species in Senegal, its northern
range was only known to extend from the region of Lake Chad to the Nile
Valley. With its occurrence in Ethiopia and Senegal its range extends across
the width of Africa, although there is still a large gap between the Senegal
and Nigerian localities.
The species is apparently rare in Ethiopia, for these are the only examples
found in spite of intensive trapping and observation at Bahadu, Koka and
other suitable localities at different seasons of the year. Because of their small
size they may elude capture by slipping through all except small mesh mist
nets, and so may be more numerous than the captures indicate.
Lonchura fringilloides, Magpie Mannikin: Two birds seen together briefly
on 24th July 1 97 1 at Didessa (9 02' N, 3 6° 09' E) were judged to be this
species. Two nets were placed in the vicinity, and caught three together later
in the day. None of this species has been seen since during several visits
extending over many weeks at different times of the year. Identification of
one bird has been confirmed by Mrs. B. P. Hall at the British Museum. The
weights of the three birds at 1400 were: 19.8, 21 .3 and 20.2 g, and all had
wing-lengths of 63 mm and were moulting their remiges.
The actual locality was on the plateau just above the east side of the
Didessa River gorge in high grass {Hyparrhenia sp.) in a large area of cleared
CombretumlTerminalia woodland. There were a very few scattered trees
remaining. Mackworth-Praed & Grant (1955) give the range of the species
in East Africa as southern Sudan, Uganda and Kenya to the Zambesi and
Zanzibar Island; its range extends westwards as far as Senegal.
Summary: The first known occurences in Ethiopia of Calidris melanotos,
Chlorocichla flavicollis, Locustella fluviatilis, Phylloscopus sibilatrix, Acrocephalus
baeticatus and Lonchura fringilloides are detailed.
Acknowledgements : I wish to extend my thanks to Dr. George E. Watson
of the Smithsonian Institution and to Dr. J. R. Schmidt of U.S. Naval
Medical Research Unit No. 3, Field Facility, Addis Ababa, Ethiopia, under
whose auspices this work was undertaken, and whom together with Mr.
K. D. Smith commented on a draft of these notes; also to Mr. Derek Good-
win and Mrs. B. P. Hall of the British Museum (Natural History), through
the courtesy of Dr. D. W. Snow, for their comments on the specimens they
examined.
References :
Brown, L. H. & Urban, E. K. 1970. Bird and mammal observations from the forests of
southwest Ethiopia. Waalia 2: 13-40.
Dowsett, R. J. & Fry, C. H. 1971. Weight losses of trans-Saharan migrants. Ibis 113:
531-533-
Ginn, P. J. G. & Brooke, R. K. 1971. The American Pectoral Sandpiper in Africa. Bull.
Brit. Orn. CI. 91: 125-126.
Kemp, A. C. 1972. A further southern African report of the American Pectoral Sandpiper.
Bull. Brit. Orn. CI. 92: 23.
Mackworth-Praed, C. W. & Grant, C. H. B. 1955. African handbook of birds. Series I, Vol. 2.
London: Longmans.
Urban, E. K. & Brown, L. H. 1971. A checklist of the birds of Ethiopia. Addis Ababa: Haile
Selassie I University Press.
Possible Female Hybrids
between Bucephala islandica and clangula
by Jon Fjeldsa
Received 2nd October 1972
Three seasons of field work at Lake Myvatn, Iceland, aroused my interest
in the status of the Common Goldeneye Bucephala clangula in that area. State-
6
ments in the literature referring to this species breeding in Iceland are un-
reliable, and based mainly on one very doubtful sight record, and one
wrongly identified specimen (Gardarsson 1968). However, in recent years
males have been found to winter regularly in south-west Iceland and may be
seen there together with unidentified females, or immature birds. It appears
further from the intensive ornithological investigation of Lake Myvatn since
i960 (Gardarsson 1968, S. A. Bengtson, pers. com., J. Fjeldsa) that some
males are present there every summer, mainly in flocks of male Barrow's
Goldeneyes Bucephala islandica. I have never seen males of the Common
Goldeneye mated to a female or defending territories in Myvatn. Neverthe-
less, I deemed it worth while to make a thorough search for female Common
Goldeneyes or hybrid specimens in the large series from Myvatn at the
Zoological Museum of the University of Copenhagen, collected mainly by
R. Hjorring and G. Dinesen at the turn of the century. All measurements
given below are from the Copenhagen collections.
At the first examination in September 1971 one female was put aside as
a possible hybrid. After having read about a hybrid male B. islandica x clangula
seen in Myvatn in 1970 by Bengtson (1972), I resumed my revision of the
collection. Among a total of 288 Goldeneyes examined, two may be hybrids.
These are a juvenile 9, Merrymeeting Bay, Maine, U.S.A. 30.10.1922 (ZMUC
69022) and an adult $, Myvatn, Iceland, 27.6.1907 (ZMUC 59275). So far
only hybrid males have been described in the literature (Snyder 1953:
Schultz 1958: Jackson 1959: Bengtson 1972), probably because possible
female hybrids will be virtually indistinguishable.
Skins of female and juvenile Common and Barrow's Goldeneyes are very
similar in general appearance although there are several slight differences,
discussed particularly by Brooks (1920). Barrow's are generally larger. In
adult females the average wing length is 215^3 mm, as against 211^4 mm
in B. clangula americana, and 199:^6 mm in B. clangula clangula. Generally, the
female Barrow's have slightly darker heads with a slight purplish lustre; the
neck is less white; the grey bar across the breast is broader; on the frontal
median wing coverts the white fringes are mainly 5 mm broad, while in the
Common Goldeneye some feathers have 8 mm long white tips. In juvenile
Barrow's the corresponding feathers are fringed with light or pale mouse
grey, not whitish. The black tipping of the greater wing coverts is an un-
reliable distinction. The most clear cut difference is the colour and the shape
of the bill. In the female Barrow's pink coloration extends backwards to
below the nostril. Seen from above the bill is more tapering. The ratio of
bill-width adjacent to the posterior angle of the nostrils, to the width, 5 mm
anterior to nostrils, averages 1.26^0.05, while in the Common Goldeneye
it averages 1.09 ±0.03. There is a slight overlap. There is a lump on the
culmen, anterior to the nostrils, although this character is shown also by
some Goldeneyes. According to Ridgway (1887) the ratio of the height of
the upper mandible at the base to the length, from loral feather edge to
anterior edge of nostrils (see Fig.) > is 1 in Barrow's, < 1 in the Common
Goldeneyes. The reliability of this character is disproved by Brooks (1920)
and in the present paper (ratio C/D in Figure). The bill nail of adult Barrow's
is 6.8 - 10.8, averaging 8.8^0.7 mm wide, rounded to form a noticeable
lump, while narrower 3.7 - 6.9, averaging 5.3±o.6 mm, more parallel-
sided and flat in Common Goldeneyes. The number of lamellae in the
mandible (which cannot be counted accurately as most anterior ones are
rudimentary) is 29-33, averaging 31.0m Barrow's, while at 33 - 37, in Com-
1.0-1
0.9-
0.8-
5
o
0.7
^oiH
a5-
oA-
Bucephala islandica
%&?
Bucephala clangula
I?
IT h
Ratio C/D
Figure
Scatter diagram for the relative height of the bill (C/D) and the relative breadth of the bill
nail (A/B) of Barrow's (solid symbols) and Common Goldeneyes (open symbols) examined.
Pulli, juveniles and adults are lumped together. It is indicated on the drawing of the Bar-
row's bill, how the measurements were taken. Two possible hybrid females are indicated
by figures i and 2.
mon Goldeneyes it averages 35.3. Gardarsson (1968, fig 1) separated the
species by plotting the width of the bill nail against the width of the bill,
adjacent to the anterior end of the nostrils. As the bill is more tapering in
Barrow's, I found the ratio of bill nail width, to the width between the
anterior angle of the nostrils, a still more reliable character. The distance
between the nostrils is similar in both species. In the diagram all specimens
examined, including pulli and juveniles, are plotted according to the above
mentioned ratio, and the ratio of Ridgway. It is indicated on the drawing of
the Barrow's bill, how the measurements are taken. Sliding callipers, ac-
curacy 0.1 mm, were used. Specimens found near the dividing line were
measured up to ten times. Specimens which are by colour of bill and plumage,
best referred to Barrow's, are plotted by solid symbols, those best referred
to Common Goldeneye, by open symbols.
The two species appear to be well separated in this way. The dots at the
top of each "dot-swarm", are mainly downy young, or juveniles not fully
grown, and deserve therefore, less attention. Only two specimens already
mentioned remain problematic by an odd combination of bill- and plumage-
characters. These are indicated by figures on the diagram.
Specimen No. i, ad. $ from Myvatn (ZMUC 59275), with brood patch'
would be placed with "islandicd" on colour and a wing length of 217 mm. It
corresponds well to other islandica females shot in midsummer. The white
edges of the median wing coverts average 6. 5 mm wide, i.e. slightly more
than in islandica; but still the closed wing gives a more black and white
variegated, less white appearance. However, the bill nail is only 6 . 8 mm
wide (ratio A/B o . 70), elevated, but with distinct parallel sides ; the bill has
not the typical tapering; ratio of bill width adjacent to posterior angle of
nostrils to the bill-width, 5 mm anterior to nostrils, is 1 . 18, which is at the
lower extreme of Barrow's. The mandible has 34 lamellae, and this is the
largest number found in specimens labelled as Barrow's.
Specimen No. 2, juv. $ from Maine (ZMUC 69022), has mainly "clangula
americana" characters; wing length 209 mm; narrower breast bar; the median
wing coverts with 4 mm wide tops of greyish white; the larger coverts with
narrow fuscous tips ; bill sides parallel, although the ratio given for tapering
is 1 . 14, which is at the upper extreme of Common Goldeneye. The nail is
6.8 mm broad (ratio A/B 0.76), and the mandible has 34 lamellae.
Among two hybrid males collected (Snyder 1953; Jackson 1959), one had
a bill corresponding to that of the Myvatn $, the other one a bill corres-
ponding to the Maine juv. 9-
Judging from these combinations of characters the above females may be
hybrids, although this cannot, of course, be stated with certainty. They have
one thing in common. They are from regions where only one Goldeneye
species is known to breed definitely, but where the other species is a regular
visitor. This applies also to the male hybrids described in the literature
(British Columbia, Seattle, New Brunswick, Myvatn). Barrow's Goldeneye
breeds mainly on the Pacific Coast and in the Rocky Mountains of Canada;
while in the coniferous zone east of the mountains it is replaced by the
Common Goldeneye; and there are very localised populations of Barrow's
only north of the timber line in northern Labrador, and maybe in the
Godthab district in West Greenland, and 800-1,000 pairs at Lake Myvatn.
No abnormal specimens were found in the series from east of the Atlantic,
where stragglers of Barrow's are extremely rare. Thus, hybrids may arise
occasionally by males forming mixed pairs in regions where they do not find
conspecific prospective mates.
Thanks are due to Mrs. F. M. Benson and to Mr. R. Wagstaffe, for kindly checking the
text of this paper, and to Dr. F. Salomonsen for critical discussion.
References :
Bengtson, S. A. 1972. An apparent hybrid between Barrow's Goldeneye Bucephala islandica
and the Common Goldeneye B. clangula in Iceland. Bull. Brit. Orn. CI. 92: 1 00-101.
Brooks, A. 1920. Notes on some American ducks. Auk 37: 353-367.
Gardarsson, A. 1968. Hvinendur a fslandi og nokkur ord um akvordun hvinendar. Nat-
turufraedingum 37: 76-92.
Jackson, M. F. 1959. A hybrid between Barrow's and Common Goldeneyes. Auk 76. 92-94.
Ridgway, R. 1887. A manual of North American Birds. Philadelphia, 631 pp., 124 plates.
Schultz, Z. M. 1958. Sight record of a hybrid male goldeneye. Murrelet 38: 11.
Snyder, L. L. 1953. An apparently hybrid goldeneye. Wilson Bull. 65 : 199.
B o
The Cape Eagle Owl Bubo capensis in Mocambiquc
by H. D. Jackson
Received 13th October 1972
When Benson & Irwin (1967) reviewed the distribution and systematics of
Bubo capensis, the species had not been recorded satisfactorily in Mocambique.
It had, however, been taken in the Inyanga highlands of Rhodesia not far
from the Mozambique border and this led Clancey (1971) to postulate its
occurrence along the entire mountainous part of the Mozambique/Rhodesia
frontier area. This frontier includes the Chimanimani Mountains, a rugged
range of quartzite and micaceous schist, and when visiting these mountains
in May 1972 I indeed heard B. capensis calling near the Mevumosi River in
Mogambique (Jackson, in press). Confirmation of its presence in the Chimani-
mani Mountains was obtained when I secured a specimen on nth August
1972 near the Mucrera River, Mozambique, at 19 53' S., 33 03' E., and
altitude 1,500 m. Two or three pairs were heard calling frequently within
5 km of my camp during the four weeks that I was there.
The specimen is an adult female taken in breeding condition, the largest
oocyte measuring 6 mm and the brood patch being devoid of feathers. Wing
(409 mm) and tail (232 mm) measurements together with the reduction of
barring on the lower abdomen and thighs show it to be B. c. mackinderi as
would be expected on distributional grounds (see Benson & Irwin, 1967).
It weighed 1,540 g (to the nearest g on an Ohaus Dial-O-Matic).
An entire leg of the Red Rock Hare Pronolagus crassicaudatus was found in
the bird's oesophagus; I am obliged to V. J. Wilson, National Museum,
Bulawayo, for confirming the identity of the prey. The enormous foot of
B. capensis suggests that it habitually preys on animals of this size; the maxi-
mum spread of the toes, measured in life from the tip of the middle claw to
the tip of the hind claw, was 103 mm. Its foot is considerably larger than that
of B. africanus and is comparable with that of the African Hawk Eagle Aquila
fascia fa. Bubo capensis is clearly a highly rapacious species.
References :
Benson, C. W. & Irwin, M. P. S. 1967. The distribution and systematics of Bubo capensis
Smith (Aves). Arnoldia {Rhodesia) 3(19).
Clancey, P. A. 197 1. A handlist of the birds of southern Mocambique. Mems Inst. Invest.
Cient. Mocamb. A, 10: 145-303; 11: 1-167.
Jackson, H. D. In press. Sight records of some birds and mammals in the central Chimani-
mani Mountains of Mocambique and Rhodesia. Durban Mus. Novit.
The breeding seasons of birds in the
Arusha National Park, Tanzania
by J. S. S. Bee sky
Received 4th October 1972
INTRODUCTION
This paper is intended to be a companion to a paper by Beesley (in press),
in which the birds and their habitats in this area were described in detail.
Breeding records were few, however, and during the thirteen months be-
tween 1st April 1971 and 30th April 1972 a great deal of effort was put into
finding nests. Of the 1,255 records in the present paper, 70% were found
during this period. It was fortunately a fairly typical period weather-wise,
and in other "normal" years the breeding seasons should be similar.
All nests recorded were seen and inspected by myself, although most of
those seen after ist April 1971 were found by members of the staff of the
Arusha National Park, and I am especially indebted to the efforts of Chacha
Mangeny, Silvesta Matacha, Sifaeli Mangure, Shauri Sandu and Anton
Wambura. For comments and advice on this paper I am grateful to Sir Hugh
Elliott and C. W. Benson.
The specific data are summarized by months in tables and the method used
by Benson (1963) has been followed, in which each record has been referred
to the month when egg-laying was calculated to have commenced. All records
are based on eggs or nestlings, except in some cases when small nidifugous
young were seen. Records of colonial breeders (marked in the tables by "C")
have been treated collectively, e.g. one colony of fifteen pairs of nesting
swifts would be set down as one record. At the foot of each table, of which
there is one for each of seven groups, the records have been converted into
percentages which are rounded off to the next whole number, i.e. 6.12
would be entered as 7. Any percentage less than five is omitted and peak
figures are shown in bold. Rainy seasons have been outlined by vertical lines
enclosing them. The nomenclature follows Mackworth-Praed & Grant
(1952-55)-
THE PHYSICAL BACKGROUND AND OTHER FACTORS
The following short description is based on Beesley (in press). The Arusha
National Park has an area of about 130 sq. km, and is situated on the eastern
side of Mt. Meru, in north-eastern Tanzania at ca. 3 15. S., 37 00' E.
Altitude ranges from 1,400 m to 4,560 m, resulting in many habitats, which
are listed below: —
1. Forest and closed woodland ranging from a dry evergreen forest at
1,450 m to a wetter montane forest up to 2,600 m.
2. Lakes and swamps, either alkaline or fresh, at 1,500 m.
3. Secondary woodlands with an understorey of scrub represented by
three main types all within an altitidunal range of 1,400 m to 1,700 m
as follows : —
a. Groves of Acacia xanthophloea.
b. Scattered remnant cedar trees Juniperus procera with an understorey
of Do done a vis cos a scrub.
c. Copses of Croton macrostachyus with an understorey of "sage-bush".
4. "Sage-bush" : A mixture of several species of soft, aromatic shrubs and
scattered trees with a ground herbage of tussock grasses which re-
places destroyed forest below 1,700 m.
5 . Heath zone at 2,300 to 3,000 m, above which is almost bare rock.
6. A diversity of small but important habitats : Streams, rain-water pools,
lake shores, grassland, forest-edge, buildings, cliffs and river/road
banks.
It should be noted that a very large proportion of the nests were found
below 1,700 m in types 3 and 4.
The climate is characterized by two rainy seasons, the short rains of
November and December and the long rains of mid-March to early June.
The short rains are generally light and occur when temperatures are tising
They are followed by the short dry season of January, February and early
March, a period which is, however, seldom rainless but is the warmest time
of the year. The long rains are heavy and temperatures begin to drop in mid-
April. June to August is the coolest time, with mist and cloud usually
occurring until late morning, sometimes accompanied by light showers.
September and October are the dryest months, although a little rain is
recorded, and temperatures begin to rise in September. At this latitude day-
light length varies minimally and its effect on birds can be discounted. The
following figures show the rainfall at a station in the Park at 1,700 m for the
year 1st May 1971 to 30th April 1972, and the mean monthly temperatures
ofArusha township which lies at 1,400 m on the southern slope of Mt.Meru: —
M J J I A S O N D J F MA
Rainfall (mm) 312 90 57 37 14 7 100 100 10 64 243 243
Temperature (C.) 19 18 17 17 19 21 21-5 21-5 21 22 21-5 21
The results of this comparatively equable climate is that the vegetation
remains green for most of the year, very few of the trees losing their leaves
and the grass drying up for only one to two months. Due to a policy of fire-
protection, fires have been very limited in number and extent for several
years. There are two periods of grass-growth with the two rainy seasons, the
main growth being during the long rains. The variety of tree species and
habitats probably results in several flushes of leaf-growth. Flowers, fruit and
cover are always available, and insects, while not appearing in a sudden,
remarkable flush with the first rains as in dryer areas, are always abundant,
more so during the rains and less so during the cold season of June to
August.
The foregoing is in contrast to some other areas in Africa, e.g. Zambia,
where the seasons are much more clearly defined, and the nesting data
presented here are therefore much more difficult to interpret. It is evident,
especially with closely related species, that detailed knowledge of food
requirements, both for parents and their young, would help explain their
breeding seasons.
BREEDING RECORDS IN TABLES BY GROUPS
The seven groupings used by Benson (1963 : 624) are followed, but not in
the same order. The Nectariniidae are included either in the Forest group or
Miscellaneous groups because of their diverse habitats and breeding seasons.
TABLE 1 WATER AND SWAMP BIRDS
Records by months
Species
Poliocephalus ruficollis
Ephippiorhynchus
senegalensis *
Hagedashia hagedash
Thalassornis leuconotus
Aythya erythrophthalma
Anas sparsa
Anas capensis
Anas punctata
Anas erythrorhynchus
Alopochen aegyptiacus
Plectropterus gambensis
Limnocorax flavirostra
Gallinula chloropus
Fultca cristata
Actophilornis africanus
Balearica regulorum
Motacilla clara
Cisticola galac totes
Totals
Percentages
No.
/
F
M
A
M
/
Jl
A
S
O
N D
recs.
rains
rains
17
1
1
2
2
4
2
1
2
1
1
2
1
6
1
1
3
1
1
1
1
1
3
2
1
1
4
1
1
2
9
1
2
1
2
2
9
1
2
1
3
3
1
1
1
2
2
3
1
2
28
11
11
3
1
1
1
5
1
2
2
3
1
2
5
2
1
1 1
104
4
4
10
17
22
10
9
9
7
6
4 2
10
17
22
10
9
9
7
6
TABLE
2 RAPTORS
2
2
2
1 1
5
1 2
1
1
19 1
3 2 5
3
2
2
1
2
1
1
1
1
1
1
32 1
5 5 7
4
2
3
3
1
1
16 16 22
13
7
10
10
Aquila rapax
Terathopius ecaudatus *
Cuncuma vocifer *
Buteo rufofuscus
Buteo oreophilus *
Falco peregrinus
Bubo lacteus
Totals
Percentages
* — all records from the same nest in different years
Water and Swamp Birds (Table i): There are no records of the Ardeidae
and few of the smaller Anatidae and Rallidae, but such as there are show a
long season of seven months during the long rains and following months
when non-permanent water-levels are still high (there are very few records
from the permanent alkaline lakes), thus agreeing with Benson (1963).
Naturally the length of the season varies with the amount of rainfall, and the
three records of Thalassornis kuconotus in October were obtained in a year of
high rainfall when the fresh- water pools remained for long at a high level.
The Rallidae, also Poliocephalus ruficollis and Actophilornis africanus, all of
whose numbers fluctuate considerably during the year and from year to year,
have some diversity in their collective breeding times depending on the rain-
fall, although P. ruficollis, which is always present, shows a definite peak from
April to August. The resident pair each of Ephippiorhynchus senegalensis and
Balearica regulorutn appear to keep to one rather short season. Moticilla clara,
whose habitat is perennial streams, has two breeding periods similar to the
insectivorous birds (Table 6) in October and February, thus differing from
most other water-birds. Cisticolagalactotes is difficult to place, perhaps because
of inadequate data.
Raptors (Table 2): Here, despite the few records, there seem to be two
seasons indicated. Aquila rapax, Terathopius ecaudatus, Cuncuma vocifer (of
which there is only one pair of each in the Park) and most of the Buteo
rufofuscus lay from March to June to rear young in the damp, cool months of
June to August, while Falco peregrinus, Buteo oreophilus and a few of the
B. rufofuscus lay from August to October and rear young in the warm, dry or
warm, wet period before and during the short rains. It is difficult to see the
advantage of laying in the earlier, wetter part of the year, as feeding nestlings
takes place while cover to conceal the prey is still thick. The prey-cover factor
does not affect C. vocifer but B. rufofuscus preys mainly on mole-rats Tachyoryctes
sp. in the Park. Perhaps these animals surface more frequently when the
soil is damp and sticky. The data for A. rapax and T. ecaudatus are equally
puzzling. There are plenty of adult grasshoppers and juveniles of grass-
nesting birds at this season, but reptilia and small mammals must be difficult
to see. Certainly these two species hunt a great deal in the dryer short-grass
plains beyond the Park borders, where A. rapax also breeds at this season.
Possibly the Park A. rapax follow the rhythmn of those extra-territorial birds.
Referring to Benson (1963) and Benson et. al (1964), in Rhodesia and Zambia
A. rapax lays in the dry season in contrast to the Park birds, but there is
agreement in T. ecaudatus laying during the rains. The owl, Bubo lacteus, fed
its nestling during September and October, when some ground cover had
dried or withered, and rising temperatures would have increased the activity
of prey.
13
Grass Birds (Table 3): The majority of the records show, as may be
expected, that nesting takes place after the short rains and during the long
rains when the grass has grown, providing nesting material, cover and food.
With a few species there is some indication of two peaks. The Cisticolas, also
Chloropeta natalensis, Prima subflava and Melocichla mentalis, are mainly long
rain breeders, but occasionally nest in the short rains.
TABLE 3 GRASS BIRDS
Records by months
J F M A M J J I A S O N D
recs.
rains
rains
2
1
: 1
11
5
5
1
4
2
2
5
2
1
1 1
15
2
2
1
5
2
3
3
1
2
16
1
3
6
2
3
1
7
2
1
2
2
4
1
2
1
5
2
2
1
8
1
1
1
1
2
1 1
14
3
1
1
1
5
3
15
1
3
1
1
1
5
1
1
1
6
1
1
1
2
1
2
1
1
12
3
1
1
5
2
5
1
1
1
1
1
8
2
1
1
1
2
'
L
3
1
1
1
4
3
1
14
1
2
2
4
5
163
14
14
20
27
30
23
10 1
I 4
8
7 5
9
9
12
17
19
14
6
5
5
Species No.
Centropus superciliosus
Chloropeta natalensis
Cisticola chiniana
Cisticola woosnami
Cisticola cantans
Cisticola brachyptera
Prinia subflava
Melocichla mentalis
Ploceus spekei C
Ploceus intermedins C
Ploceus castaneiceps C
Hyphanturgus ocularis
Othyphantes reichenowi
Amblysopi^a albifrons C
Euplectes capensis
Coliuspasser laticauda
Spermestes cucullatus
Spermestes nigriceps
Lagonosticta rubricata
Coccopygia melanotis
Estrilda astrild
Totals
Percentages
Numida mitrata
Francolinus hildebranti
Francolinus squamatus
Burhinus capensis
Charadrius pecuarius
Charadrius tricollaris
Stephanibyx coronatus
Hoplopterus armatus
Caprimulgus fraenatus *
Caprimulgus poliocephalus
Caprimulgus fossil
Mirafra africana
Ant bus si mi lis
Macronyx aurantiigula
Saxicola torquata
Totals
Percentages
* — This name follows White (1965)
Of the Ploceidae, Ambljospi^a albifrons and Coliuspasser laticauda begin to
nest in the short dry season and continue until the end of the long rains, but
Ploceus spekei, P. intermedius and Euplectes capensis have one short season,
almost entirely during the long rains. Ploceus spekei and P. intermedius are
14
TABLE 4
GROUND
BIRDS
6 3
1 1
1
2
1
1
12 1
2
5
2
2
3 1
1
1
13 2 1
1
2
4
1
2
6 1 1
1 1
2
2
1
1
9
2
6 1
8 1 2
2
1
1
1
5
2
3
1
1
4 1
1 1 1
13 2 2
3 2
1
2
1
7 2 2
1
1
1
18 3 2
2
4
4
3
109 11 16
8 4 5
9 4
6
17
12
10
7
11 14
7 5
9
6
16
12
10
7
conspicuous colonial nesters, and any nesting at other times would not have
been missed. Perhaps as with Quelea spp. the young are fed much on young
grass seeds, or the type of grass used for building does not attain sufficient
length until late in the rains. Ploceus castaneiceps, Hyphanturgus ocularis and
Othyphantes reichenowi probably breed at any time of the year, contrasting
markedly with the other Ploceidae. Estrilda astrild breeds almost entirely in
the long rains, and Coccopygia melanotis follows at the end of the rains in
rather cool conditions.
Ground Birds (Table 4) : The two dryest periods of the year, January-
February and September-October, serve as the main breeding seasons for
these birds as a whole, although probably due to the porous, quick-draining
soil of the Park, some breeding takes place even during the rains. Some
species apparently have only one breeding season, and others, for example the
Caprimulgidae, have two, both avoiding the rains. Hoplopterus armatus is
noteworthy, laying in the cool damp month of June to rear young in cold
July. Mirafra africana resembles the grass-birds in breeding during the long
rains, thus agreeing with Benson (1963). However, Macronyx aurantiigula
records differ from the Macronyx species of further south (Benson et al. 1964)
in nesting mainly outside the rains. Macronyx aurantiigula and Mirafra africana
both occupy similar habitats in the same areas of the Park, bush-scattered
and/or tussock grassland. Saxicola torquata has an extended season of six
months when, it would appear, two broods are raised. Evidently it can breed
in the short rains, but not the long rains.
TABLE 5 FOREST BIRDS
Records by months
Species
Tympanistria tympanistria
Tauraco hartlaubi
Halcyon albiventris
Melittophagus oreobates
Tockus alboterminatus
Lybius melanopterus
Buccanodon leucotis
Ari^elocichla nigriceps
Ari^elocichla milanjensis *
Alseonax adustus
Tchitrea viridis
T urdus olivaceus
Cossypha semirufa
Seicercus umbrovirens
Psalidoprocne holomelaena
Dryoscopus cubla
Pholia sharpii
Onychognathus walleri
Stilbospar kenricki
Zosterops eurycricotus
Nectarinia taca^e
Cinnyris mediocria
Cyanomitra olivacea
Nigrita canicapilla
Serinus burtoni *
Totals
No.
/
F
M
A
M
/ fl
A
S
N D
recs.
rains
rains
1
1
8
1
1
1
1
1
2 1
6
1
1
1
2
1
4
2
2
1
1
12
1
1
1
1
1
2
2
3
20
3
2
1
2
9
3
3
1
1
1
2
1
1
6
2
1
3
13
2
1
1
3
2 4
3
1
1
1
2
1
1
1
1
5
1
1
2
1
2
1
1
1
1
2
2
12
1
1
2
1
1
5
1
2
1
1
4
2
2
4
1
2
1
5
1
2
1
1
4
2
1
1
2
2
125
11
7
5
6
5
1 3
7
21
30
17 12
9
6
5
6
17
24
14 10
Percentages
- Records based on obesrvation of food and nesting material being varried
!5
Forest Birds (Table 5): The breeding season of the majority of this type
agree with comments by Moreau (1950) and Benson (1963) in that there is
one fairly well-defined season avoiding the heavy rains. Nesting begins in
September when the forest is comparatively dry and the temperature starts
to rise. Maximum laying is in October, which is warmer, and most young
are being fed on the flush of insects during the rains of No vember and December.
There are a few apparent exceptions. Alseonax adustus and Tchitrea viridis have
a second season in January, February and March, and Cyanomitra olivacea has
its only season at this time.
TABLE 6 INSECT AND FRUIT EATING BIRDS OF
WOODLAND AND SECONDARY BUSH
Species
No.
Records by months
F M A M J J I A S O N D
Eurystomus glaucurus
10
3
4
3
Halcyon chelicuti
5
1
1
2
1
Melittophagus bullockoides C
6
1 1
2
2
Melittophagus pusillus
9
1 1
1
4
2
Phoeniculus purpureus
3
1
2
Rhinopomastus cyanomelas
2
1
1
Colius striatus
72
12
15
9
14
1
1
3
4
11
2
Lybius leucocephalus
2
2
Tricholaema lacbrymosum
48
8
12
11
1
2
1 1
2
3
6
1
Campethera nubica
5
1
1 2
1
Campethera abingoni
2
1
1
Dendropicos fuscescens
1
1
Thripias namaquus
3
1
1
1
Mesopicos goertae
3
2
1
Motacilla aguimp
4
1
1
1
1
Pycnonotus barbatus
155
33
38
19
8
12
1 7
16
19
2
Dioptrornis fischeri
33
12
3
1
3
5
7
1
1
Batis molitor
8
2
1
2
3
Apalis flavida
25
1
1
5
1
2 6
8
1
Phyllolais pulchella *
2
1
1
Sylvietta whytii
10
1
1
2
2
2
2
Camaroptera brevicaudata
4
3
1
Campephaga sulphur at a
3
1
2
Nilaus afer
1
1
Lanius collaris
25
4
3
5
1
2
1
4
3
2
Laniarius aethiopicus
7
1
2
1
1
2
Tchagra senegala
4
1
1
1
1
Tchagra aus trails
14
3
5
1
2
1
2
Chlorophoneus sulfureopectus
2
1
1
Par us fringlllinus
3
1
2
Orlolus larvatus
6
1
1
1
2
1
Onychognathus morlo
6
2
1
2
1
Buphagus africanus
2
2
Buphagus erythrorhynchus
7
2
1
2
2
Totals
492
83
86
56
33
24
10 6
5 47
63
62
17
Percentages
17
18
12
7
5
10
13
13
* _
- Nest-bui
lding observe
d only
Insect and Fruit-eating Birds of Woodland and Bush (Table 6) : The two
seasons are exemplified well here, one before and during the first part of the
short rains and the other in the hot, short dry season and beginning of the
16
long rains. Of the birds for which there are sufficient records, 60% have two
seasons and 36% have one main season with a few nestings at other times.
Only one bird, the migrant Eurystomus glaucurus, has one definite season. It
arrives in August and leaves in January-February. No other species in this
table is known to be migratory.
Species with the most records have their maximum breeding during the
second period, i.e. in the short dry spell preceding the long rains, but a few
show preference for the first period, i.e. before and during the early part of
the short rains. In the case of Pycnonotus barbatus there is an indication of a
slight recrudescence of nesting in May, but this may relate to a second
attempt by nest-predated birds.
It is interesting that Thripias namaquus starts nesting in May, thus avoiding
competition with other Picidae. The two species of Meropidae begin during
the cold damp months of June and July. Also it is surprising that Apalis
fiavida has its second peak in May, rearing young in the cool month of June.
TABLE 7 MISCELLANEOUS GROUPS
Records by months
Species
Columba arquatrix
Streptopelia semitorquata
Streptopelia Ingens
Streptopelia capicola
Stigmatopelia senegalensis
Treron australis
Agapornis fischeri
Apus caffer
Apus niansae C
Apus horus C
Hirundo abyssinica
Hirundo senegalensis
Hirundo smithii
Pytonoprogne fuligula
Riparia plaudicola C
Serinus flavivertex
Serinus striolatus
Carduelis citrinelloides
Nectarinia kilimensis
Chalcomitra amethystina
Chalcomitra senegalensis
Anthreptes collar is
Cinnyris venustus
Totals
Percentages
No.
/
F
M
A
M
/
//
A
S
AT D
recs.
rains
rains
4
1
3
39
2
5
5
2
4
3
1
1
3 8
4 1
1
1
2
1
1
1
1
9
1
3
1
3 1
7
3
2
2
1
1
3
3
5
3
2
3
1
2
7
1
1
2
1
2
19
2
1
4
2
1
2
2 5
9
3
1
2
1 2
7
2
5
4
1
3
15
5
3
1
2
1
1
1
17
4
3
3
3
2 2
9
1
1
4
1
1
1
18
2
3
1
2 4
3 3
14
1
1
2
2
2
1 2
2 1
13
2
1
1
1
1
2
3
2
44
9
10
2
4
1
1
9
4
4
251
28
31
20
31
24
17
18
8
10 31
21 12
12
13
8
13
10
7
7
13
9 5
, Miscellaneous Groups (Table 7) : Of the Columbidae there are only three
species, all mainly arboreal, with sufficient records for interpretation. Treron
australis has one season with apparently two peaks, one in November and
another in February. Streptopelia semitorquata nests throughout the year as
further south (Benson et al. 1964), but with peaks in October and February-
March, months which produce abundant fruit, insects and seeds,
c 17
Discounting Apus caffer for which there is but one record, only two species
of Apodidae have been found nesting in the Park. Apus horus and A. niansae
both breed in April at the height of the long rains in high river-banks, which
is puzzling and agrees with Benson's remarks (1963) {A. niansae also breeds
in cliffs higher up the mountain). As these birds are colonial nesters it is
hardly possible that they were overlooked at other times of the year. Both
these species feed a good deal over the lakes, and perhaps there is a greater
availablity of food over these waters at this time. A. horus bred again, at least
during two years, in November when insect-life is known to be abundant.
The Hirundinidae show diversity in their seasons. Ptyonoprogne fuligula
has two, similar to birds of group 6. Riparia paludicola, like the Apodidae,
breeds in river-banks (but lower ones) during the long rains when there is
danger of collapsing banks, contrary to the records in Benson (1963). Hirundo
smithii, generally using buildings, nests in any month (one nest was used for
four broods in one year) and H. senegalensis, either taking over nests of H.
smithii or building in trees, has the same tendency. The colder season does
not deter these birds from nesting, so evidently insect-life is still fairly
abundant.
The Fringillidae are puzzling, as in Zambia and Malawi. Serinus flavivertex
descends from the higher levels of Mt. Meru to breed among scattered
juniper trees at 1,500 m during the wet cool months of May and June.
Carduelis citrinelloides has two seasons, each during a rainy season, one of
which is synchronous with S. flavivertex. Serinus striolatus begins with its
peak in January and continues in a lesser degree for the following six months.
C. citrinelloides and S. striolatus overlap somewhat and occupy the same
habitat (sage-bush), but the latter tends to nest lower down in the shrubs and
in areas where there are more grass tussocks.
The five species of Nectariniidae collectively breed in every month and
there is some overlapping of the peaks. The main area of overlap is in
October and November when three species show much activity. Two of
them, Anthreptes collaris and Chalcomitra amethystina, feed and nest mainly in
forest-edge and woodland. Feeding appears to take place on two levels, that
of the former in the lower storeys and of the latter in the higher storeys. The
third, Chalcomitra senegalensis, feeds and nests to a greater extent in the low
secondary sage-bush. Nectarinia kilimensis and Cinnyris venustus also breed in
this latter biotope, but N. kilimensis shows a peak in April and C. venustus two
peaks, one in January and February and another in July, both being seasons
when several components of the sage-bush are in flower.
CLUTCH SIZES
Clutch size data from the period 1st April 1971 to 30th April 1972 have
been extracted for some species, which are listed below. Comapring these
with the figures in Benson et al. (1964), which are for Rhodesia only {pp. cit. :
32), where records are sufficient for this purpose, some clutch sizes are in
agreement, i.e. in the Columbidae, Caprimulgidae, Batis molitor, Cisticola
cantans, Prinia subflava, Tchagra australis y Chlorophoneus sulfureopectus and
Nectariniidae. Others differ in that the Park's birds have smaller clutch sizes,
usually by one egg, i.e. in Colius striatus, Anthus similis, Macronyx spp.,
Pjcnonotis barbatus, Tchitrea viridis, Saxicola torquata, Lanius collaris, and
Laniarius aethiopicus.
Number of Records
C/l
C/2
C/3
Columba arquatrix
3
Streptopelia semitorquata
7
17
Caprimttlgus fraenatus *
1
7
Caprimulgus poliocepbalus
5
Caprimulgus fossii
1
Colius striatus
4
28
16
Ant bus si mi lis
2
9
Macronyx aurantiigula
2
2
Pycnonotus barbatus
10
101
5
Dioptrornis fiscberi
6
1
Batis molitor
4
Tchitrea viridis
6
Saxicola torquata
5
7
Melocicbla mentalis
1
4
Apalis flavida
2
11
2
Sylvietta wbytii
5
Cisticola woosnami
2
Cisticola cantans
2
2
6
Pr/nia subflava
2
1
4
Ptyonoprogne fuligula
1
5
Lanius collaris
2
3
Laniarius aetbiopicus
4
Tchagra australis
3
5
J
Chloropboneus sulfureopectus
2
Nectarinia kilimensis
3
Cinnyris venustus
3
24
3
Anthreptes collaris
3
3
C ardeuelis citrinelloides
2
2
Serinm striolatus
9
2
C/4
This name follows White (1965)
SUMMARY
The Arusha National Park lies within a regime of two rainy seasons, the
short rains occurring in November and December, and the long rains from
mid-March to early June. The result is two main flushes of plant growth and
insect-life. The dry seasons are short, cloud cover being prolonged during
the longest of them, and the effect of drought on the vegetation is minimal.
These conditions enable some breeding to take place throughout the year,
and make records difficult to interpret. The breeding records are divided into
seven categories and are listed in tables as follows : —
i . Water and swamp birds show much diversity but have a maximum of
breeding activity during and after the long rains.
2. Raptors divide into two groups, one nesting early, during the long
rains, the other later, in the ensuing dry season. Buteo rufofuscus, the
most numerous, nests throughout both periods.
3. Grass birds show some diversity, but some species (i.e. Cisticolas and
Ploceidae) appear to start with the short rains, continuing with little
break to the end of the long rains. However, some Ploceidae can
breed at any time of the year.
4. Ground birds usually nest during the two dry periods, but most species
have only one breeding season.
5 . Forest birds generally have one season, from September to December,
before and during the short rains.
6. In insect and fruit-eating birds of woodland and secondary bush, two
breeding seasons show up well, one before and during the first part
of the short rains, and the second in the hot, short, dry season and
beginning of the long rains, 60% of the species having two seasons.
J 9
7. Miscellaneous groups show a considerable diversity, but nesting
generally avoids the cold month of August and, surprisingly, the more
congenial month of September.
An appendix of clutch sizes is included, and some comparisons made with
Rhodesian data.
References :
Beesley, J. S. S. In press. The birds of the Arusha National Park. Journ. E. Afr. Nat. Hist.
Soc. <& Nat. Mus.
Benson, C. W. 1963. The breeding seasons of birds in the Rhodesias and Nyasaland. Proc.
i$th Internat. Orn. Congr. II: 623-639.
Benson, C. W., Brooke, R. K. & Vernon, C. J. 1964. Bird breeding data for Rhodesias and
Nyasaland. Occ. Nat. Mus. S. Rhod. 27B: 30-105.
Mackworth-Praed, C. W. & Grant, C. H. B. 1952-55. African handbook oj birds. Series I,
Vols. 1 & 2. London: Longmans.
Moreau, R. E. 1950. The breeding seasons of African birds — 1. Land birds. Ibis 92: 223-267.
White, C. M. N. 1965. A revised checklist of African non-passerine birds. Lusaka: Government
Printer.
On the supposed genus Vetrochelidon
by Allan R. Phillips
Received 8th November 1972
Brooke (1972: 55-56) maintains Petrochelidon apart from Hirundo because it
"has a red rump and a virtually square tail and does not build an entrance
tunnel to its mud pellet nests : some species are colonial breeders". (He also
removes from Hirundo certain other species, placing them in Pseudhirundo and
Cecropis, with which I am not here concerned, for lack of field experience.)
The nests of the common swallows are well known, and publicized in a
multitude of short articles and local and general works, besides the classic
Bent "Life Histories" (Bent 1942). Contrary to Brooke's claim, the type
species of "Petrochelidon", "P. pyrrhonota" or a/bifrons, the Cliff Swallow,
does quite uniformly build an entrance tunnel to its retort-shaped nests. The
Cave Swallow, "P." fulva, usually does not, but its nesting is quite variable;
see for example Wetmore and Swales, 1931 : 320-322.
Tail shape is quite variable among species generally conceded, even by
Brooke, to belong to Hirundo, sensu strictu. I cannot regard rump colour as a
good generic character; and since none of Brooke's other characters hold
true, I continue to regard "Petrochelidon" as inseparable from Hirundo, as
previously (Phillips, Marshall, and Monson, 1964).
Of more importance, perhaps, is the nasal operculum (cf. Ridgway, 1 904 ;
Brodkorb, 1968); but its variation (presence or absence) cuts right across the
colour and nesting variations, which may prove more basic.
References :
Bent, A. C. 1942. Life Histories of North American Flycatchers, Larks, Swallows and their
Allies. U.S. Nat. Mus. Bull. 179.
Brodkorb, P. 1968. Part Five. Birds, p. 260-451. {In W. F. Blair et at, Vertebrates of the
United States. Second Edition.) New York, etc. : McGraw Hill Book Co.
Brooke, R. K. 1972. Generic limits in old world Apodidae and Hirundinidae. Bull. Brit.
Orn. CI. 92: 53-57.
Phillips, A., Marshall, J. & Monson, G. 1964. The Birds of Arizona. Tucson: Univ. Ariz.
Press.
Ridgway, R. 1904. The Birds of North and Middle America . . . Part III. U.S. Nat. Mus.
Bull. 50, pt. III.
Wetmore, A., & Swales, B. H. 1931. The Birds of Haiti and the Dominican Republic.
U.S. Nat. Mus. Bull. 155.
Nesting of the Roseate Tern (Sterna dougallii)
near Dar es Salaam
by D. Keith Thomas & Hugh F. I. Elliott
Received 16th October 19 j 2
The current interest in seabirds of the East African coast has prompted us
to dig out some long unpublished notes and photographs of a Roseate Tern
colony which established itself near Dar es Salaam from July to September
i960. This was recorded soon afterwards by one of us (Thomas 1962), but
with few details. Otherwise, there seem to be no records of the breeding of
this species in what is now Tanzania, except for those quoted by Mackworth-
Praed & Grant (1952), namely Zanzibar (in October: a breeding male and a
chick marked "Chumbe Zanzibar 3/10/27" are in the British Museum
(Natural History) collection); coral reef near Pemba (June and July); Kisite
islands near Tanga (July); and, "almost certainly", Mafia (Aug. and Spet.).
The Dar es Salaam colony was discovered by D. K. Thomas on 7th August
i960, on an undercut ("mushroom") coral islet of about 50 mm circumfer-
ence and 4.5 m in height, one of several strung out along an extensive reef
which more or less dries out at low spring tides. This reef, the Hammand,
lies to the north of Outer Makatumbe ("Lighthouse") island, some 4 km
from the harbour entrance. The number of adult terns was estimated at
about 300 and this was confirmed when we both visited the site on 21st
August. On that occasion six to ten pairs of terns were also found to be
nesting on a smaller islet about 100 m away. Some eggs had already been
hatched on 7th August, suggesting that the colony had been occupied by
mid- July. However, some eggs were also still being incubated on 21st
August and, on our final visit on 3rd September, when only 20-30 adults
remained in the vicinity, a few unfledged chicks were being fed.
The flat-looking but in fact very rough and deeply creviced tops of the
islets, both of which were accessible only at fortnightly intervals (in the case
of the main colony thanks to the collapse of a large segment of the overhang),
were covered by a dense tangle of vegetation, identified (L. H. Brown in
litt.) as mainly composed of Salvadorapersica and Cissus quadrangularis^ although
one baobab had managed to obtain a ledgement. The eggs had been laid on
the bare coral or, occasionally, on a sparse mat of plant detritus, which looked
as if it had been brought together as nest material. There was particularly
good cover for young birds on the dense vegetation and crevices. In most
cases only a single egg had been laid, but three C/2 were noted. Seven eggs
measured averaged 43.5 x 31.5 mm (extremes 46.5 X 30.5, 40.5 x 31
and 45 x 32); egg colour was very variable, from pale bluish white almost
unmarked through stone-grey with fine black and ashy speckling to deep
buffy with large blotches of dark brown and lavender-grey. There was a
frequent tendency for the markings to form a central band.
Observations of this colony and examination of the two specimens col-
lected, an adult and chick, now in the British Museum, support the conclusion
that tropical populations of Roseates tend to be smaller, whiter below (with
no discernible pink flesh) and redder-billed when in breeding condition and,
in juveniles, with the pale grey of mantle and scapulars more heavily overlaid
with buffy mottling, compared with populations of the sub-tropics or
temperate regions whether to north or south. Our birds' bills looked red with
a black tip rather than black with red at the gape, although in the adult
specimen collected only 20. 5 of the 39 mm of the upper mandible was in fact
red, or just over half, This specimen most closely matches those from Ceylon
in the British Museum collections, which are attributed to the subspecies
korustes originally described from the Andamans. But specific nomenclature
Roseate Tern colony on one of two coral islets on Hammand reef, Dar es Salaam. Top : the
islet at low spring tide: broken segment on left allowed access to colony — about 50 terns
visible in photograph; part of Outer Makatumbe island in right background. Middle: the
mat of Cissus quadrangular is and Salvadora persica on the summit — looking west over outer
anchorage of Dar es Salaam towards city. Bottom: egg on coral rag in the Salvadora) Cissus
tangle, a few leaves and twigs having apparently been. assembled as nest material.
is an unsatisfactory way of recording the intergradations of this wide-ranging
species, especially as all the characters that have been used except size (where
the variation is only of the order of five or six per cent) are invalid unless the
birds compared are known to have been at the same stage in the breeding
cycle.
References :
Mackworth-Praed, C. W. & Grant, C. H. B. 1952. African handbook of birds. Series I, Vol. 1.
London: Longmans.
Thomas, D. K. 1962. Birds — further notes on breeding in Tanganyika. Tang. Notes &c Recs.
58/59: 200.
The Relationships of Picathartes
by Charles G. Sibley
Received 18 th October 1972
The systematic position of the West African genus Picathartes has been
examined by several authors in recent years. Sclater (1930) and others before
him had placed Picathartes in the Corvidae but Lowe (1938) proposed that it
is a member of the Sturnidae. Amadon (1943) suggested that Picathartes may
be an "aberrant member of the thrush-babbler assemblage" and Delacour
(1950) placed it in a special tribe, Picathartini, in the Timaliinae. Delacour &
Amadon (195 1) reviewed the problem and the evidence and concluded "that
Picathartes has far more in common with babblers than with starlings, crows,
or other passerine families". Serle (1952) reviewed the nestling behaviour,
eggs and juvenile plumage of Picathartes and found nothing inconsistent with
a relationship to the Corvidae. He noted that the babbler Eupetes of south-east
Asia, Indonesia and Borneo resembles Picathdrtes in general proportions,
texture of the plumage and bare skin areas. Although Serle advocated com-
parisons with all passerine groups he thought that Picathartes should be kept
"in or near the Corvidae". Bannerman (195 1 ; 1953) circumvented the prob-
lem by placing Picathartes in its own monotypic family but indicated his
acceptance of Lowe's position by placing it next to the Sturnidae. White
(1962) retained Picathartes in the Corvidae "largely because no convincing
reasons for moving it have yet been given" (p. 4). Deignan (1964: 442)
recognized a subfamily. Picathartinae, near the Timaliinae in the Musci-
capidae. Hall & Moreau (1962) included Picathartes in the Timaliidae,
Moreau (1966: 161) referred to "the babblers Picathartes gymnocephalus Jonas"
and Hall & Moreau (1970: 149) "tentatively" followed the arrangement
presented by Deignan {he. cit.).
In a recent study (Sibley 1970) the egg-white proteins of Picathartes
gymnocephalus were compared with those of many other passerine groups
using the technique of starch gel electrophoresis. I found (p. 62) that the
patterns produced by Picathartes egg white were more like those of the
timaliids than those of coivids or sturnids, and thus supported the conclusions of
Delacour & Amadon (195.1). But this evidence fell short of providing proof
of such a relationship because of the limitations of the starch gel technique.
I am now able to report on additional studies of the egg-white proteins of
Picathartes using the electrophoretic method of isoelectric focusing in acryla-
mide gel (IFAG). This techinque is described in Sibley & Frelin (1972).
With IFAG many more proteins can be resolved and the resulting patterns
provide an improved basis for systematic comparisons. I have discussed the
use of electrophoretic data in systematics in previous papers (e.g. Sibley
1970).
Two sets of comparisons, using pH ranges of 3-10 and 6-4, were carried
out. That Picathartes is a babbler, rather than a sturnid or a corvid, is easily
established but it is even clear that Picathartes is especially close to the African
timaliine genus Turdoides. Some 21 protein bands can be seen in the^H 3-10
gels and all of them are identical or nearly so between Picathartes gymno-
cephalus and Turdoides jardineii and T. ieucopygius. The other babblers, for which
egg-white specimens were available, namely, Trichastoma sepiarium, T. bicolor,
Pomatostomus temporalis and P. superciliosus, basically agree with Picathartes
and Turdoides, but show some minor differences in the positions or concen-
trations of some proteins.
In the "magnified" patterns of the pH 6-4 region the detailed similarities
between Picathartes and Turdoides are striking. Ten proteins can be resolved
in this limited range and for six of them Picathartes and Turdoides have identi-
cal isoelectric points. The other four show small differences in isoelectric
points of a magnitude I have often observed between congeneric species. In
fact, the two species of Turdoides are only slightly more alike than Picathartes
and either of the Turdoides The most striking contrast is found in the com-
parisons between Picathartes and the Malayan Trichastoma and the Australian
Pomatostomus. Both show greater differences when compared to Picathartes
than those noted between Picathartes and the African Turdoides.
It thus seems clear that Picathartes is a babbler and that its closest living
relatives are probably the members of the genus Turdoides. It will be necessary
to compare the egg-white proteins of Picathartes with those of all genera of
African babblers before we can be absolutely certain that Turdoides is closest
to it. However, of the babblers occurring in Africa Turdoides seems to be the
most likely close relative of Picathartes. The resemblances between Picathartes
and Eupetes noted by Serle (1952) are almost certainly the result of con-
vergence but should be investigated further. The egg white of Eupetes is not
available but it, and that of all species, would be welcomed by the writer.
In view of this new evidence I recommend that Picathartes be placed next
to Turdoides and that they be arranged no farther apart than as tribes in the
same subfamily. In my opinion they are closely related genera and the only
reason for separating them in different tribes is to recognize their superficial
morphological differences.
AC KNOWLEDGEMENTS
The first egg white of Picathartes which I received came to me through
the kindness of Mr. J. T. Menzies, at that time resident in Sierra Leone. The
specimen used in this study was collected in Ghana by Dr. L. G. Grimes to
whom I am also grateful for a memorable trip to see Picathartes gymnocephalus
in the field during a visit to Ghana in 1964. The other egg white specimens
mentioned were provided through the kindness of Dr. David R. Wells, Mr.
Tony E. Bush and Mr. Gordon B. Ragless. Mr. Jon E. Ahlquist gave
generously of his help and advice and technical assistance was provided by
Miss Dorothy J. Moore. The National Science Foundation supported the
study under grant GB-6192X.
References :
Amadori, D. 1943. The genera of starlings and their relationships. Amer. Mus. Novit. 1247:
1-16.
Bannerman, D. A. 195 1. The birds of tropical West Africa. Vol. 8. London: Crown Agents.
— 1953. The birds of west and equatorial Africa. Vol. 2. Edinburgh & London: Oliver &
Boyd.
24
Deignan, H. G. 1964. Subfamily Picathartinae, p. 442 in Checklist of birds of the world, vol. 10,
ed. E. Mayr & R. A. Paynter, Jr., Cambridge, Mass.: Museum of Comparative
Zoology.
Delacour, J. 1950. Les timaliines. Additions et modifications. L'Oiseau et R.F.O. 20:
186-191.
Delacour, J. & Amadon, D. 195 1. The systematic position oiPicathartes. Ibis. 93 : 60-62
Hall, B. P. & Moreau, R. E. 1962. A study of the rare birds of Africa. Bull. Brit. Mus. {Nat.
Hist.) 8: 315-378.
— 1970. An atlas of speciation in African passerine birds. Brit. Mus. (Nat. Hist.) publ. no. 68o,
pp. xv & 423.
Lowe, P. R. 1938. Some anatomical and other notes on the systematic position of the genus
Picathartes, together with some remarks on the families Sturnidae and Eulabetidae. Ibis 80:
254-269.
Moreau, R. E. 1966. The bird faunas of Africa and its related islands. New York & London:
Academic Press.
Sclater, W. L. 1930. Sy sterna avium ethiopicarum. London: Brit. Orn. Union.
Serle, W. 1952. The affinities of the genus Picathartes Lesson. Bull. Brit. Orn. CI. 72: 2-6.
Sibley, C. G. 1970. A comparative study of the egg-white proteins of passerine birds.
Bull. Peabody Mus. Nat. Hist. 32: 1-131.
Sibley, C. G. & Frelin, C. 1972. The egg white protein evidence for ratite affinities. Ibis 114:
377-3 8 7-
White, C. M. N. 1962. A revised check list of African shrikes, orioles, etc. Lusaka: Government
Printer.
Blood parasites from a Red-chested Owlet
Glaucidium tephronotum
by John E. Cooper
Received 6th November 1972
The Red-chested Owlet Glaucidium tephronotum Sharpe is a little known forest
species ranging from Ghana east to Kakamega forest, Western Kenya (White
1965 ; Ripley & Bond 1971). There is a paucity of data on the biology of the
species (Brown 1970) and I have been unable to trace any references to its
diseases or parasites. In this note I report the finding of three species of
blood parasite in a captive specimen of the Zaire and Uganda race, G. t.
medje Chapin.
Full details of the bird's history have been impossible to obtain. Dr. C.
Pennycuick received it in late 1969 from Dr. M. P. C. Fogden, from Western
Uganda; Pennycuick used the bird in his wind-tunnel experiments in Nairobi
for some time until it was noted, on 5 th July 1970, to be depressed in
appearance; no treatment was given and it was found dead the next day.
A full post-mortem examination was carried out at the Veterinary Research
Laboratory, Kabete, Kenya on the day of death. The bird was an adult male,
thin (weight 67 g) with some damage and soiling of feathers but carried no
ectoparasites. The skin is now in the National Museum, Nairobi. There was
no internal fat and the liver was congested. The gizzard contained only
chitinous arthropod remains. Two small nematodes were found on the
abdominal air sac walls and these were later identified by the British Museum
(Natural History) as a Demsidocercella species. No parasites were found in the
intestinal tract but a faceal sample showed three Capillaria eggs, several
unidentified spiruroid or filaroid eggs and a few coccidial oocysts. Culture of
intestine yielded Escherichia co/i, but heart blood was bacteriologically sterile.
Routine blood smears from the heart of the bird were fixed with methyl
alcohol and stained with Giemsa's stain. Examination revealed three blood
parasites — the Protozoa Haemoproteus and Leucocyto^oon (parasitizing the red
25
and white blood cells respectively) and numerous microfilariae (Nematoda)
within the plasma. It was not possible to identify these parasites further from
a blood smear. There were no haematological abnormalities and no evidence
of anaemia. Both blood Protozoa and microfilariae have been recorded
previously from owls but as Keymer (1972) and Cooper (1972) point out,
there is little data on their pathogenicity. In the present case there was no
evidence to suggest that they were associated with disease and the cause of
death remains obscure. In addition to the Red-chested Owlet above I have
examined blood from 12 owls in Kenya (eight Tyto alba^ one Ciccaba wood-
ford/t, one As/'o capensis, one Bubo africanus and one Bubo lacteus) but in none
of these have I found either Protozoa or microfilariae. It is, of course, possible
that the parasites in the Red-chested Owlet were acquired in captivity but all
three parasites are vector (arthropod) borne and it would seem more likely
for the bird to become infected in its natural habitat in the foiest than in
captivity in the laboratory. Whatever the source however, the presence of
three blood parasites in the blood of one specimen of Glaucidium tephronotum
medje would appear to warrant record.
I am grateful to Dr. C. Pennycuick for data on the owlet and for permission
to record these findings, to Dr. I. F. Keymer for confirming the identification
of the blood parasites and to Mr. G. C. Backhurst for valuable comments and
advice on this paper.
References :
Brown, L. H. 1970. African Birds of Prey. London: Collins.
Cooper, J. E. 1972. Veterinary Aspects of Captive Birds of Prey (In press).
Keymer, I. F. 1972. Diseases of Birds of Prey. Vet. Rec. 90: 579-594.
Ripley, S. D. & Bond, G. M. 1971. Systematic Notes on a Collection of Birds from Kenya
Smithsonian Contributions to Zoology number 1 1 1 .
White, C. M. N. 1965. A revised Checklist of African Non-Passerine Birds. Lusaka: The
Government Printer.
Notes on the birds of Rangiroa, Tuamotu Archipelago,
and the surrounding ocean
by D. T. Holy oak
Received 20th November 1972
While travelling between Tahiti and the Marquesas Islands in 1972 I twice
passed through the northern end of the Tuamotu Archipelago, and spent two
days on Rangiroa atoll. Very little information is available on seabirds at sea
in this region and the land-birds are little known, so it seems worthwhile
recording the limited observations I was able to make.
On 1 3 th August I was on the French Navy frigate "L'Enseigne de Vaisseau
Re Henri" about 30 miles east of Makatea at dawn. I was able to make
almost continuous observations as the ship passed between Rangiroa and
Arutua, then between Ahe and Manihi, reaching a position just north of
Manihi by dusk. In September I returned from the Marquesas Islands on the
American yacht "Sea Star", reaching n point about mid-way between the
Marquesas and Tuamotu Islands by dawn on 3rd September (12 19' S,
143 57' W.), spending the next two days sailing to Rangiroa, than 6th-7th
September on the north side of Rangiroa atoll.
Rangiroa is one of the largest atolls in the world, being about 45 miles
from east to west and 20 miles from north to south. It consists of a ring of
coral and coral sand, mainly between 100-300 m wide, reaching a width of
600 m in a few places, and broken by a few small sea gaps. The shallow
lagoon enclosed is so wide that the other side of the atoll cannot be seen in
many places. The land rises to a maximum of about 5 m above sea-level and
supports a meagre vegetation dominated by Scaevola bushes and coconut
palms. Near the small villages there are numerous introduced trees and
bushes. The population is about 800, nearly all of them Polynesians who live
by fishing, copra production and subsistence farming.
The northern Tuamotus are within the latitudes of the Southeast trade
winds, which blow with little variation in strength and direction. The ocean
falls away to great depths close to the islands, and is affected by the easterly
South Equatorial Current, which moves up to about five miles a day near
Rangiroa. Water disturbances as this current reaches the steep shores of the
archipelago probably account for the rich marine life that supports the large
seabird populations.
Pterodroma rostrata, Tahiti Petrel : Birds identified as this species were seen
at 12 20' S, 144 oo' W (one, 3rd September), 12 30' S, 144 35' W (one, 4th
September) and 12 50' S, 145 ° 23' W (five, 5th September). At least 20 other
gadfly petrels seen were either this species or the similar P. alba. The seven
listed above were all distinguished from that species at close range by their
larger size, heavier bill, heavier less buoyant flight and the absence of a small
light patch under the chin. Under the variable light conditions at sea I found
it difficult to detect the differences in colour of the upperparts of these two
species. These are the first records for rostrata in Tuamotus waters, although
it breeds in both the Society and Marquesas Islands (King 1967).
Pterodroma alba, Phoenix Petrel: One seen at 12 46' S, 145 ° 19' W on 4th
September was the only one positively identified, although some of the
unidentified gadfly petrels mentioned under rostrata were probably this
species; it breeds in the Tuamotus.
Pterodroma arminjoniana heraldica, Herald Petrel: Only one seen, at 12 19' S,
143 57' W, on 3rd September; breeds in the Tuamotus.
Pterodroma neglecta, Kermadec Petrel: Two seen on 5th September (13 43'
S, 1 47 10' W and 13 50' S, 147 17' W) were distinguished from the rather
similar white phase of P. arminjoniana heraldica by the dull brown underwings
with an oval white patch restricted to the area around the base of the pri-
maries (not a longer, diffuse, light stripe), slightly larger size, and the way
in which they remained lower over the water when banking. The only other
records of this species in the Tuamotus are three specimens collected by the
Whitney Expedition that are now in the American Museum of Natural
History.
Pterodroma leucoptera, White-winged Petrel: Two seen together on 3rd
September at about 12 20' S, 145 ° 21' W, are the first records for the
Tuamotus area. They were seen down to a range of about 1 5 m and were
noted at the time as, "small gadfly petrels, grey above with dark 'W' back
back pattern, blackish patch on crown extending to lower nape, all-white
underparts, small bill, underwings virtually all-white with no trace of a dark
line extending inwards from the alula, flight quick with rapid wing-beats".
About ten small and medium-sized gadfly petrels were not seen well
enough for species identification.
Puffinus pacifus, Wedge-tailed Shearwater: About ten were seen at intervals
on 13th August; one seen about ten miles from the north coast of Rangiroa
on 9th September, and about ten birds that were probably of this species
seen at 13 43' S, 147 10' W, at dawn on 5 th September. All were of the dark
colour morph. Breeds in the Tuamotus.
27
Puffinus nativitatis, Christmas Shearwater: One seen at about 12 46' S,
145 ° 19' W, on 4th September; breeds in the Tuamotus.
Nesofregetta albigularis, White-throated Storm Petrel: One seen 13th
August about 50 miles W.N.W. of Makatea, and another on 5th September
at about 13 02' S, 146 20' W. Both were light phase birds, and they appear
to be the only records for the Tuamotus other than old ones of "dark storm
petrels" quoted by King (1967), which could have been other species of
Hydrobatidae or Bulweria. Identification was certain as the white throat patch
bordered below with black, white belly and butterfly-like flight were noted
with both birds.
Phaethon aethereus, Red-billed Tropicbird: An adult seen at about 1 3 45 ' S,
1 47 1 1' W, on 5 th September. This is the first record for the Tuamotus area.
Identification was certain as the bird repeatedly flew around the mast of the
yacht, giving clear views of a scarlet bill, long white central tail feathers and
narrow black bars on the back.
An unidentified immature tropicbird was seen about 40 miles north of
Rangiroa on 5th September; both P. Upturns and P. rubricauda breed in the
Tuamotus (King).
Sula sulci, Red-footed Booby: A total of several hundred was seen close
to the islands, singly and in small groups. None were seen more than about
30 miles from land except for a single bird at about 12 28' S, 144 15' Won
3rd September. Three out of 231 birds counted were of the light phase;
breeds in the Tuamotus.
Sula leucogaster, Brown Booby: Single birds were seen frequently close to
the islands, a total of about 20 being counted; none were seen more than 30
miles from land. Breeds in the Tuamotus.
Fregata minor, Great Frigatebird : The only bird definitely identified as this
species was an adult male seen on 6th September about two miles north of
Rangiroa. Breeds in the Tuamotus.
Fregata ariel, Lesser Frigatebird : Apparently commoner than minor; 1 1 were
identified on or near Rangiroa and one near Ahe, most of them soaring over
the atolls. A further 20 or so unidentified and immature frigatebirds were
seen near to the islands, and single unidentified immatures were also seen far
from land on 2nd September (around 12 22' S, 144 05' W) and 3rd Septem-
ber (12 25' S, 144 35' W). Breeds in the Tuamotus.
Egretta sacra, Reef Heron: Common on Rangiroa. They fed mainly on fish
in the clear, shallow water along the shore, but also in a small grassy marsh
and on bare areas of the atoll among scattered bushes where insects appeared
to be the main prey. I will comment elsewhere on counts of the two colour
morphs made on Rangiroa and other islands in the Tuamotus (Holyoak ms a).
G alius gallus, Domestic Fowl: Numerous around houses on Rangiroa, but
no feral populations exist.
Pluvialis dominica, Lesser Golden Plover: Common along the seaward shore
of Rangiroa and on grassy areas of the atoll, about 30 being seen altogether.
Most birds were moulting from summer to winter plumage in early Septem-
ber, but the stage of moult varied considerably between individuals.
Prosobonia (Aechmorhynchus) cancellatus, Tuamotu Sandpiper : One was seen
on a bare area of Rangiroa about three miles south-east of the airstrip, on 6th
September. It was alone, and feeding on insects taken from the ground in a
bare area of coral among scattered herbaceous plants. For the five minutes I
was able to watch it was absurdly tame, allowing me to approach to within
about 2 m before running further away; throughout this time it gave a soft
"pew" "pew" squeaking call almost continually. After I had followed it for
28
about 40 m it suddenly flew (disclosing heavy moult of the primaries and
secondaries) and continued low over the beach until lost to view.
This species is endemic to the Tuamotu Archipelago, where it breeds only
on small atolls having no men, cats or rats {mss by Quayle & Beck in American
Museum of Natural History). As atolls like this are gradually becoming
scarcer its continued survival is in doubt unless some small atolls are set aside
as reserves. Presumably the bird I saw was merely a casual visitor to Rangiroa
as it has not been recorded there before and this island is unsuitable for
breeding owing to the abundance of men and rats.
Tringa incana, Wandering Tattler: Numerous along the seaward shore of
Rangiroa, about 20 being seen in total. Most of them were still in summer
plumage.
Sterna {Thalasseus) bergi, Crested Tern: About ten seen close to the shores
of Rangiroa on 6th~7th September. These were adults accompanied by one
or two flying juveniles each. An adult was seen to fly towards a juvenile that
was following and giving squeaking calls, then to pass a small fish to its bill
about 20 m up above open water. Another juvenile was watched standing on
a beach attempting to swallow a reef fish about 8 cm long, 5 cm deep and 1
cm thick; eventually it gave up and left the fish on the beach.
Sterna fuscata, Sooty Tern: Numerous around the islands, hundreds being
seen around Rangiroa on 6th~7th September. Both days on Rangiroa flocks
of up to 60 birds were frequently seen flying high along the atoll to the south-
east, calling frequently. Only three in juvenile plumage were seen despite
careful inspection of "noddies". Up to five daily were also seen far from land
on 13th August and 3rd-4th September; breeds in the Tuamotus.
Sterna iunata, Grey-backed Tern: Two seen at about 12 46' S, 145 ° 19' W
on 4th September, and six near the shore of Rangiroa on 6th September;
breeds in the Tuamotus.
Procelsterna cerulea, Blue-grey Noddy: At least 50 seen close to shore of
Rangiroa on 6th~7th September; none seen away from land. Appeared to
be breeding in coral blocks along shore of Rangiroa, as sereval birds flew out
from beneath them as I passed and it is known to breed elsewhere in the
Tuamotus (King).
Anous stolidus, Brown Noddy: Large numbers seen whenever I was within
20 miles of land, feeding flocks seen off Rangiroa containing up to about a
thousand birds of this species. Apparently less numerous than A. tenuirostris.
Breeds commonly in coconut palms on Rangiroa and elsewhere in the
Tuamotus. I saw several well-grown young birds in coconut palms and
adults were seen to fly from them. Few seen far from land, my only records
being of five at about 13 43' S, 147 10' W on 5 th September and five more
at about 13 52' S, 147 50' W the same day.
Anous tenuirostris, Black Noddy: Large numbers seen whenever I was
within about 20 miles of land, but none further out. Several feeding flocks
seen near Rangiroa had several thousand birds of this species, and flocks of
scores or hundreds were always in view from the shore. Nests abundantly in
coconut palms on Rangiroa, where I saw numerous nestlings at all stages of
growth.
One was watched flying back and forth alone over lagoon water only 5-8
cm deep, periodically dipping in to water to catch a tiny fish in its bill and
touching the bottom with its feet, then resting there briefly with opened
wings half-raised before flying up and starting to hunt again. A Reef Heron
feeding in the water lunged at this bird several times when it passed close by.
29
Gygis alba, White Tern : Common near to land, many hundreds being seen
feeding close to Rangkoa, where they were breeding in coconut palms and
other trees (two small young seen). Far from land small numbers were seen
daily, maximum 16 on 2nd September.
This species is conspicuous in the large feeding flocks of terns characteristic
of the Tuamotus as it flies to and fro about 10-25 m above the water,
periodically dipping to catch small fish. Anous stolidus and A. tenuirostris
characteristically fly beneath the White Terns, usually only 1-5 m above the
water. All three species feed throughout the day in large numbers ; the White
Tern also feeds at night, but I obtained no proof that the Anous species do so
too.
In the bright sunlight usual in this area the white flashes as Gygis turns to
dip to the water surface are conspicuous at a great distance, and are used by
local fishermen to locate fish shoals. Presumably the white colouration also
enables other Gygis and other species of fish eating-bird to spot fish shoals
easily. When solitary birds or small flocks of this species first start to feed they
give a long, high-pitched "weeee" squeaking call almost continually. This
must make them conspicuous to birds that have not found food from several
miles away, as from the near-silence of a yacht under sail I often heard them
before seeing them at ranges of at least 1 km; I did not notice this call when
travelling on motor powered ships and boats, probably because of the engine
noise. Perhaps the call aids in the co-ordination of nocturnal feeding, as I
frequently heard it at sea during the night.
In view of the number of White Terns I saw far from land between the
Marquesas and Tuamotu Groups it seems likely that the endemic Marquesan
subspecies microrhyncha occurs in Tuamotu waters, although records are
lacking.
Ptilinopus (purpuratus) coralensis, Tuamotu Fruit Dove: Seven or eight seen
in tall scrub and low trees in a coconut plantation in the village on the north
side of Rangiroa. I did not see any in the scrub and coconut plantations away
from the village where there were few trees other than coconuts. They were
comparatively tame, often not flying away until I walked in full view of the
bird to within about 4 m. It moves freely from about 1 2 m up in the tree-tops
to the ground, feeding on the ground and in very low bushes as well as in
taller bushes and trees. Its movements through foliage are agile, with long
hops between branches, rapid sideways runs down them and hanging head
downwards to reach food. The general shape and outline of this form are
distinctly slimmer and more passerine-like than those of the related ptilinopus
in the Society Islands. Its flight is direct, but not very rapid, with quick,
regular, rather shallow wing-beats that make a whistling sound.
I saw them eat black berries about 7 mm in diameter from a low bush, small
green flowers of a low tree, and many insects pecked from the foliage of
bushes and the ground with a quick darting action. Bruner (1972 p. 75) also
records them eating insects on the ground, and grass seed.
The usual song is similar to that of Society Island Ptilinopus, a soft "0000-
()()()o-( )ooo-ooo-ooo-oo-oo-o-o-o-o-o" accelerating towards the end as it
dies away. An alternative song similar to those heard in the Society Islands
was also heard several times and tape-recorded, "ooa-00 oooa-00 oooa-00
oooa-oo". They frequently sang from low in bushes as well as high perches.
After singing one bird was seen to fly down in to a low bush and perch
near to another about 3 m up. It then walked towards the other bird, allo-
preened its crown and nape for about a minute (without huddling close to the
30
bird being preened), stepped back and preened itself. Head-scratching in this
species is direct (foot lifted straight to head).
A local woman showed me a nearly fully-built nest less than a metre above
the ground in a patch of thick, low scrub among taller bushes in a coconut
plantation. The nest was well-hidden from above, and consisted of a flimsy,
flat platform of thin twigs and slender pieces of vine stem. Local people all
agreed that it lays a single pale brown egg in a flimsy, flat nest. The eggs of all
Ptilinopus so far known are white, except that superbus lays white or creamy
white eggs (Goodwin 1967); thus my informants may have been wrong
about the egg colour. I was told that it also nests higher up in bushes but that
nests within about 1 5 cm of the ground are common.
Goodwin (/. c.) and others have lumped this form with Ptilinopus purpuratus
of the Society Islands, along with chalcurus from Makatea and chrysogaster from
the Leeward Society Islands. It is always difficult to decide what taxonomic
status should be afforded island forms that replace each other geographically,
and any treatment is little better than an informed guess. Generally the best
basis from which to guess would seem to be the extent of differences in
closely related forms that achieve sympatry. InPtilinopus the sympatric species
mercierii and dupetithouarsi of the Marquesas Islands do not differ more in
plumage than some of the forms included in purpuratus, and they appear to
differ less in behaviour and ecology than do purpuratus and coralensis, (Holy-
oak in prep, and ms b).
P. purpuratus purpuratus and P. p. chrysogaster of the Society Islands have
very similar general behaviour and ecology (Holyoak ms b), and differ from
coralensis in being almost entirely arboreal, eating fewer insects, nesting
higher in trees and in general body shape. It seems unlikely that either would
survive for very long in the habitat of the other. P. p. chalcurus is intermediate
in plumage between coralensis and nominate purpuratus, but differs from both
in its darker mauve cap. The biology of chalcurus is practically unknown,
but the raised atoll of Makatea probably offers conditions that are inter-
mediate in many respects between those of the mountainous Society Islands
and the low atolls of the Tuamotus. As Makatea is also between the Society
Islands and Tuamotus geographically, it seems likely that chalcurus does
represent a link between coralensis and the nominate form. Thus it may be
best to keep all of those forms together in one species despite the marked
differences in their general ecology; however, any course of action is largely
! arbitrary.
Acrocephalus atypha, Tuamotu Warbler : Three birds were seen in an area of
tall bushes under scattered coconut palms near the airstrip on Rangiroa. They
were very tame, approaching through open branches to within 2 m of me in
a very bold manner. In general their actions when moving through vege-
tation were similar to those of Acrocephalus arundinaceus ; although their flight
appeared weaker.
They pecked small insects from twigs and leaves as they hopped rapidly
up and down branches, and one was seen to catch a small insect in flight.
Calls were similar to those of A. caffra of the Society and Marquesas Islands,
a low "chru" being frequently repeated. The song was also similar in struc-
ture to that of caffra, but less continuous, less rich and less varied. Typically
the song consisted of the "chru" call followed by a short whistling phrase,
e.g. "chru-peee-tu-tu-pee-wee" followed by a pause before the song was
repeated with a different arrangement of whistling notes. They sing at all
levels from high in coconut trees to low down in bushes, and appeared to
sing most vigorously around sunset.
3 1
Singing birds replied vigorously and approached me with raised crown
feathers and half-opened wings when I played tape recordings of their song
to them. The whistling songs of a variety of unrelated American birds, in-
cluding wrens, thrushes and a parulid, evoked equally vigorous responses.
Presumably as A., atypha is the only songbird in the Tuamotus it has lost the
ability to differ entiate between songs of conspecifics and other species that
related birds of continental areas have. It is also likely that the present varied
and musical song has not degenerated, solely because there is a need for it
in intraspecific relations.
I saw several old nests built in bushes from 4-8 m above the ground,
usually among trailing vines. One I examined was a substantial, deep, cup-
shaped structure built of grass stems, long leaves and the fine stems of vines,
with a lining of slender vine stems. It measured about 12 cm from top to
bottom and was built in a fork of the branch of a bush among a thick vine
growth.
AC KNOWLEDGEMENTS
My visit to Polynesia was made possible by a fellowship of the Winston
Churchill Memorial Trust, and preparatory work at the American Museum
of Natural History was financed by a grant from the Frank M. Chapman
Memorial Fund. I am also grateful to the officers and crew of "L'Enseigne
de Vaisseau Re Henri" and to J. Cunha, Mr. & Mrs. J. P. Kavanaugh, and
Mr. & Mrs. R. Pozzi on the "Sea Star" for hospitality and assistance.
Mr. E. Bryan jr., Mile. K. Dubois, Dr. C. Jouanin, Dr. B. Salvat, Dr. D. W.
Snow and M. C. Tissier also assisted me in various ways, and I am grateful
to Dr. W. R. P. Bourne and Mr. W. B. King for commenting on my manu-
script.
References :
Bruner, P. L. 1972. Birds of French Polynesia. Honolulu: Bishop Museum.
Goodwin, D. 1967. Pigeons and doves of the World. London: British Museum (Natural
History).
Holyoak, D. T. ms a. Significance of colour dimorphism in Polynesian populations of
Egret ta sacra.
— ms b. The birds of the Society Islands.
— in preparation. The birds of the Marquesas Islands.
King, W. B. 1967. Seabirds of the tropical Pacific Ocean. Preliminary Smithsonian Identification
Manual. Washington: Smithsonian Institution.
First Report of Pitta e. erythrogaster from Leyte
by Kenneth W. Prescott
Received 24th November 1972
Some time ago Dr. Kenneth C. Parkes questioned me concerning a pitta
which I had collected on 4th July 1945 . Although the recorded locality of the
specimen was Leyte, which attracted Parkes' attention, it was taken aboard
a ship which was anchored a few miles off the coast of central Leyte in the
Leyte Gulf. McGregor & Worcester (1906: 68) listed twenty-three Philippine
Islands from which P. e. erythrogaster has been recorded. Leyte was not one of
these although it was recorded from the neighbouring islands of Samar to
the north, Mindanao to the south, Cebu and Panay to the west and south-
west, and Masbate to the north-west. In discussing the zoologically distinct
island groups of the Phillippines, McGregor & Worcester (1906: 4-5)
grouped Samar, Leyte and Bohol together, placed Cebu in a class by itself,
regarded Mindanao and the immediately adjacent islands as by themselves,
and Panay and Masbate were placed with the central Philippine group.
Although in 1906 McGregor & Worcester did not record this pitta from
Leyte or Bohol, subsequently McGregor (1909: 414) listed it from Bohol
but not from Leyte. To date, strangely enough, it has not been reported
from Leyte. A glance at the map discloses the fairly large Leyte literally
encircled by islands on which P. e. erythrogaster is found; moreover, the
zoologically related Samar is contiguous with Leyte to the north-east. My
specimen, although collected offshore, rather strongly suggests that P. e.
erythrogaster exists on Leyte.
At dusk the bird flew aboard the well lighted U.S.S. Jamestown, A.G.P.-3,
through an open porthole alighting on a table in a crew's mess where it was
quickly pounced upon by a boatswains-mate whom I had frequently taken
on collecting trips. He brought the still living bird to me which I subsequently
made into a skin, now UMMZ-113, 932 (sex, female?) in the University of
Michigan, Museum of Zoology collection. The Jamestown was one of
several hundred lighted ships anchored that evening in Leyte Gulf. That the
pitta chanced my way was a stroke of good luck, although we were anchored
much closer to shore, two-four miles, than the other larger war ships.
For this specimen I am grateful to the quick thinking George, BM/ic,
and to the U.S. Navy which allowed me to collect birds while serving (then
Executive Officer of the Jamestown) during a time of war. I am indebted to
Dr. Parkes who called my attention to the importance of the locality in which
the pitta was collected.
References :
McGregor, Richard C. & Worcester, Dean C. 1906. A Hand-List of the Birds of the Philip-
pine Islands. Department of the Interior, Bureau of Government Laboratories ; No. 36.
January. Manila.
McGregor, Richard C. 1 909. A Manual of Philippine Birds. Part II. Bureau of Sciences,
Manila.
Supplementary Notes on the Birds of
Point Tombo, Argentina
by Jeffery Bos wall
Received jth December 1972
In company with Donaldo Maclver and Douglas Fisher I returned to Point
Tombo, Argentina, from ioth-26th October 1972 to complete a BBC
television film on the biology of the Magellanic Penguin Spheniscus magellani-
cus. The visit was earlier in the season than that in 1971, the ornithological
results of which were published by Boswall and Prytherch (1972). A few
supplementary notes are now set out with emphasis on the seabird species
known to nest. Not all bird species seen are mentioned.
The notice prohibiting access to Point Tombo in the interests of preserv-
ing the ecology of the area proved to be meaningless. Between 2,500 and
3,000 people visited the area in the austral spring and summer of 197 1-2,
mainly in December, January and February, and mainly at weekends, They
wander at will and in our experience are particularly attracted to the part of
the penguin colony without bushes, where the birds are most conspicuous
and where the burrows cave in very readily. Also, we saw that Kelp Gulls
Larus dominicanus follow in the wake of humans and carry off penguin eggs
left unguarded. The uncontrolled wanderings of tourists is a cause of great
concern to local conservationists.
33
Eudyptes crestatus y Rockhopper Penguin ; Four dead birds on the shore.
Spheniscus magellankus, Magellanic Penguin: Specially detailed notes were
kept on this bird to be used as a basis for the film commentary. Also a
separate paper is being prepared. However it is important to tidy up here
some points made in the earlier Point Tombo paper. The phrase "peak of
laying" in paragraph two under S. magellankus should read "peak of clutch
completion". Point Tombo is now known not to be "the northernmost
colony of this species on the Atlantic coast apart from 50 pairs reputed to
nest at Punta Leon . . . (64 30' W. 43 ° 05' S.)". Juan Munoz (in discussion,
1972) knows of a colony at Punta Clara (65 12' W. 43 57' S.). Murphy is
mis-quoted in the earlier paper; his only evidence that this species nests
"From the Gulf of San Matias, Argentina, southward" was not Magellan's
sighting. Murphy also refers to Burmeister, who wrote that "from about
latitude 41 ° S. on the Patagonian coast the Jackass Penguins are more or less
regular denizens of the islets, peninsulas and mainland beaches". The fact that
Durnford (1 878) found no penguins at Point Tombo at the end of December
surprised Boswall & Prytherch, but in 1972 a new fact came to light. It is
that in about 1930, when Carlos La Regina of Estancia La Perla was in his
early teens, only a very small part of the area now occupied by nesting
penguins was in use by them. Since then they have spread greatly.
With regard to breeding season, it is worth mentioning here, since the
first paper was particularly concerned with this subject, that the peak of
clutch completion in 1972 was not as early as 7th October, as Boswall &
Prytherch speculated, assuming the birds lay eggs at the same time every
year. It was well after 16th October and before 21st October. On nth,
inspection of 100 nests showed 52 with birds but no eggs, 35 with one egg
and 13 with two eggs. Comparable figures for 16th were 21,31 and 48; and
for 2 1 st October, 8, 17 and 74 plus one nest with three eggs. It must be con-
cluded either that the incubation period of the closely related species S.
demersus is not the same as that of magellankus, or that egg-laying at Point
Tombo was differently timed in 1971 and 1972. 1971/72 was an exceptionally
cold winter in Patagonia and locals advised us that, at the time of our arrival,
temperatures were still much below the usual ones for the time of year. But
whether the coming ashore of the penguins would be influenced by local
weather is not known, except that Carrara (1952) says "... the arrival on
land for nesting is from the first days of September until the beginning of
October; but there are small variations every year. The climatological con-
ditions, specially the temperature, play a most important role in the dates of
arrival on land". Meyer de Schauensee (1970) says of distribution of S.
magellankus "Breeds . . . coast of Argentina north to Santa Cruz". In fact the
species nests north to the next province, Chubut.
Diomedea melanophris, Black-browed Albatross : One seen in flight.
Macronectes giganteus, Giant Fulmar (or Petrel): A single bird was seen at a
floating penguin carcase, and on another day a group of about six birds were
similarly engaged. Conway (1971) saw Giant Fulmars attack apparently adult
penguins at Point Tombo.
Fulmarus glacialoides, Southern (or Silver-grey) Fulmar: One seen just off-
shore on 22nd October.
Phalacrocorax magellanicus, Rock Cormorant; P. bouganivillii ', Guanay Cor-
morant; and P. albiventer, King Cormorant. All three species were back at
the mixed colony .The two last were just starting to collect nesting material
on nth October. According to Carlos La Regina (personal discussion, 1972),
34
cormorant guano was taken from Point Tombo in about 1965 by a Japanese
company.
Rodolpho Escalante (pers. comm. 1972) has drawn my attention to the
fact that the observations by Erize (1972) on the first Atlantic-coast nesting
of P. bougainvillii had first been published by Navas (1970). Erize (op. cit,)
quoted Meyer de Schauensee (1970) to the effect that the nearest nesting
Guanays, measured by the shortest sea route, were nearly 3,000 kms away on
Mocha Island in Chile. Johnson (1965), however, says that this island, which
lies at Lat. 37 30' S, was deserted by 1962 and that only the colony known to
him in Chile is on Pupuya Islet at Lat. 3 3 58' S. about 400 kms further to the
north.
Boswall & Prytherch (1972) quote Olrog (1968) on P. albiventer breeding
on "the inland lakes in the province of Neuquen". Escalante (1971) quotes
Navas {Neotropica 1970 (16): 140-144) to the effect that the birds in Neuquen
are Blue-eyed Cormorants P. atriceps, and not P. albiventer. Since the latter
"has been much confused with one or more races of P. atriceps" (Murphy
1936), it is worth mentioning that a colour photograph of those at Point
Tombo was published by Erize (1970) and that two colour transparencies of
mine are deposited in the University Museum of Zoology, Cambridge.
Phoenicopterus chilensis, Chilean Flamingo : Two parties of about a dozen
birds seen. One party appeared to be feeding in the sea.
Cygnus melancoryphus, Black-necked Swans: Seven in flight on 22nd,
twelve on 23rd.
Tachyeres patachonicus, Flying Steamer-duck: The identification of this
species was confirmed by close field observation and the wing length of a
drake found dead. Also, the identification of the birds in Parrinder's photo-
graph as flightless Flying Steamer-ducks has been further confirmed by
Milton Weller and O. S. Pettingill (pers. comm., 1972): "The birds are
evidently moulting, being temporarily without most of their remiges, except
for their tertials". The sizes of six eggs found in a nest just above high water
mark on 22nd October were: 84.8 x 55 .8 mm, 83.2 x 54.6 mm, 83.0 x
54.2 mm, 85.0 x 5 5-6 mm, 83.1 x 56.1 mm., and 84.6 x 55.5 mm. The
respective weights were 140, 130, 130, 145, 140 and 135 grammes. A bird was
put off this nest, but otherwise the birds were always in pairs, the two
keeping very close together. As in the previous year, the pairs were spread
out along the coast and each pair could always be found approximately where
expected, (see also Prytherch, Boswall & Maclver, in preparation).
Vanellus chilensis, Southern Lapwing: One pair present and rather noisy.
Charadrius falklandicus, Two-banded Plover: One very young chick seen
on 13 th October.
Thinocorus rumicivorus, Least Seedsnipe: J. Daciuk tells me he has seen
chicks of this species at Point Tombo.
Chionis alba, Snowy Sheathbill : Small parties near the extremity of the
point; others in the penguin colony tidying up the remains of penguin eggs
left by gulls or skuas.
Catharacta Skua, Great Skua: Skuas were in pairs and back in their breed-
ing areas of the previous year. They would fly over to inspect any human
intruder but did not mob. One was seen to snatch a penguin egg from a nest
left unguarded because of human disturbance. Others were active in the
burrow area of the penguin colony robbing Kelp Gulls of penguin eggs.
With the exception of one buff-streaked bird, all had very dark plumage.
Leucophams scoresbii, Dolphin Gull : Up to seven together in the penguin
colony trying to "cash in" on penguin eggs stolen by Kelp Gulls.
35
Lams dominicanus, Kelp Gull : Already in occupation of their nesting areas
and very clamorous at the intrusion of an observer. A number of Kelp Gulls
were very active in the penguin colony taking eggs, but they frequently lost
them to skuas.
Sterna himndinacea, South American Tern : Up to ten birds presumed to be
of this species were seen on most days.
Sterna eurygnatha, Cayenne Tern : Four birds presumed to be of this species
seen on 13 th October.
Upucerthia dumetaria, Scale-throated Earthcreeper : A nest which had been
exposed on 23rd by a human was brought to my attention. The tunnel had
been 1 . 9 metres long, and had been hollowed from a vertical bank of soil-
cum-shingle. At the end was a scant nest of dried grasses and five smooth
white eggs. At least one bird returned and the same day tried to "mine"
further beyond the nest, but no more activity was seen.
Lessonia rufa, Rufous-backed Negrito: A female was put off a nest built at
the base of a small plant growing among rocks near the shore. It contained
three eggs on 23rd, two eggs and one chick on 24th, and three chicks on
25 th.
Diuca diuca, Common Diuca Finch : Birds almost certainly of this species
were taking grass into a hole a couple of metres from the exposed Scale-
throated Earthcreeper nest and it was presumed to be an unused burrow of
that species.
AC KNOWI EDGEMENTS
I am grateful to Robin Prytherch for checking the paper through, and to
Judy Billington for typing it.
References :
Boswall, Jeffery & Prytherch, Robin J. 1972. Some notes on the Birds of Point Tombo,
Argentina. Bull. Brit. Orn. CI. 92 (5): 11 8- 129.
Carrara, I. S. 1952. Lobos marinos, pinguinos y guaneros del littoral maritimo e islas adyacentes
de la Republic a Argentina. La Plata.
Conway, William. 1971. Predation on Penguins at Punta Tombo. Animal Kingdom August
issue 1971 : 2-8.
Durnford, Henry. 1878. Notes on the birds of Central Patagonia. Ibis (4) 2: 389-406.
Erize, Francisco. 1970. Cormorants of South America. Animals 13 (5): 235-237.
Escalante, Rodolfo. 1971. Addenda and Corrigenda to Aves Marinas del Rio de la Plata, ip.
Montvideo: Barriero and Ramos.
Johnson, A. W. 1965. The Birds of Chile. Buenos Aires: Graficos.
Meyer de Schauensee, R. 1970. A Guide to the Birds of South America. Narbeth, Penn:
Livingston.
Murphy, R. C. 1936. The Oceanic Birds of South America. New York: American Museum of
Natural History.
Navas, Jorge R. 1970. Nuevos registros de Aves para la Patagonia. Neotropica 16 (49):
11-16.
Prytherch, R. J., Boswall, J., & Maclver, D. In preparation. Casual Notes on the Flying
Steamer-duck.
Regular Occurrence of Porphyrula martinica in
South Africa
by W. Roy Siegfried & P. G. H. Frost
Received 2jth November 1972
Ohlson (1972, Bull. Brit. Orn. CI. 92: 92-93) has reported the occurrence of
the American Gallinule Porphyrula martinica on the mid Atlantic islands of
St. Helena and Ascension. The species is also a regular visitor to Tristan da
Cunha (Elliott, 1957, Ibis 99: 545-586), To date. 11 specimens have been
36
obtained in South Africa (not 12 as stated by Broekhuysen 1971, Ostrich 42:
78-79). These birds were found within 160 km of Cape Town (33° 55' S,
1 8° 22' E). This paper reports a further record of P. martinica from Cape
Town, and draws attention to the apparent regularity with which the species
occurs in South Africa.
The bird was found at Camps Bay, 10 km from Cape Town, on 20th May
1972. It was rescued in an exhausted state from a cat, but rapidly recuperated
when fed raw, lean beef. The bird was a first-year male in the process of
acquiring adult plumage. It weighed 160 gm, 63 % of the mean weight for 20
£$ recorded by Hartman (1961, Smith. Misc. Coll. 143: 1-91). Other re-
corded weights of immatures (unsexed) recovered in South Africa are even
lower, < 150 gm and 137 gm (Rowan & Winterbottom 1963, Ostrich 34:
249-250; Winterbottom 1965, Ostrich 36: 90-91). Mensural data taken from
the bird are: culmen and shield 45-5 mm, wing 183-0 mm; tarsus 64- 5 mm.
The primaries and secondaries were unworn. The tail feathers were moulting.
The adult purple abdominal and breast feathers were still sheathed at their
bases, but the new bronze feathers of the upper parts were unsheathed.
This represents the twelfth record of P. martinica in South Africa. Two
birds were recovered in April, four in May, four in June, one in August and
one of unknown date. During the decade 1 962-1 972, nine birds (all im-
mature) appeared in South Africa. 1968, 1969 and 1971 were the only years
in which American Gallinules were not recorded. Thus, the species may be
regarded as a more or less annual vagrant to the southern tip of Africa.
The prevailing winds over the south Atlantic during the austral autumn
and winter are predominantly westerly, with winds of gale force being
common. These conditions would favour forced down wind crossings of the
south Atlantic by American Gallinules blown off course while either migrat-
ing or dispersing along the south eastern seaboard of the new world. It
is somewhat remarkable how regular and often this relatively weak flying
species makes the 8000 km journey to South Africa.
The peak of occurrence in South Africa (May-June) is about a month
later than the peak on Tristan da Cunha (March-May, Rowan & Elliott, C.
pers. comm.). This could suggest that the birds occurring in South Africa
are secondarily derived from those on Tristan, where, having at least partially
recovered, and still possibly being affected by the urge to migrate, they
leave the Tristan group only to be transported westerly again to the southern
tip of Africa. This suggested movement may explain why, in spite of the
number of records down the years, there have been few reports of adults
and no reports of attempted breeding on Tristan.
This may also explain the species' occurrence on Ascension and St.
Helena. Ohlson {pp. cit.) suggested that the Gallinules recorded there could
have arrived via the south easterly trade winds (the prevailing winds in the
mid Atlantic) after having been driven to the proximity of the African coast
by the west winds of the south Atlantic. He alternatively suggested that the
American Gallinules may have been derived directly from South America by
being transported along the upper west wind layer that blows across the mid
Atlantic at altitudes of 500 m and higher. However, in as much as they can
be taken to mean anything, the dates of occurrence (in June) of the species
on these islands slightly favour the first suggestion.
The above comments are, of course, largely speculative. The lack of data
on weights of birds occurring in any of these areas precludes any interpre-
tation of movements based on the relative weight losses of individuals sur-
viving the south Atlantic crossing. In future, it might be possible to examine
37
stomach contents, specifically grit, and feather minerals (see Hanson & Jones
1968, Biol. Notes Illinois Nat. Hist. Surv. 60) in an attempt to determine not
only an individual's origin but possibly also any stop-over localities. Anyone
likely to gain possession of American Gallinules on either the mid or south
Atlantic islands or in South Africa should bear this in mind.
The Nesting of Feral Pigeons Columba livia in Trees
by Jejfery Bos wall
Received 1 9th January 1973
At Derek Goodwin's suggestion I made a few notes when in Buenos Aires,
Argentina, on 30th October 1972, on the nesting of Feral Pigeons Columba
livia in the trees of the city's sidewalks and plazas. Plaza San Martin, for
example, has a large number of very mature trees of a species that fork at
anything from about three metres to about nine metres from the ground. In
the crooks of these, the birds commonly build their nest, see Plates 1 and 2.
Photo: Jejfery Boswall
Plate i To show tree nest-site of Feral Pigeon Columbia livia, Plaza San Martin, Buenos
Aires, Argentina. The nest is situated in the major "crook" of the tree immediately
above the observer's head.
38
Photo: Jeffery Bo snail
Plate 2 Feral pigeon Columba livia on typically exposed tree nest-site, Plaza San Martin,
Buenos Aires, Argentina.
Apart from the branches of the trees, the nests were fully exposed to the sky.
One particularly exposed nest was built on the stump of a sawn off branch.
The nest material was mainly small dry twigs. The half-dozen nests I could
see into had small young. The birds also nest commonly on city buildings, I
was told, and as in other cities are commonly fed by the human populace.
One (rather poor) colour photograph submitted to Goodwin for deter-
mination of the colour pattern types brought the following comment, "there
seem to be 16 barred blues (the wild colour-pattern), 23 dark blue chequers,
velvets or blacks (probably all these but hard to say . . .). one dominant red
(ash red in American parlance), one dark blue chequer with many white
primaries, and one that is either a mealy or a light drizzle". (
Tringa hypoleucos Linnaeus breeding in East Africa
by G. R. Cunningham <& van Someren
Received i$th December 1972
The East African Natural History Society have been considering the question
of naturalisation of some species generally regarded as migrants or other
introduced species to East Africa. Forbes-Watson (1972).
39
The writer proposed that Himantopus himantopus (Linneaus) and Recurvi-
rostra avosetta Linnaeus, which breed regularly at Lake Magadi Kenya, should
be included, along with T. hypoleucos, as birds are present throughout the
year and have been recorded as breeding, van Somerens (191 1).
Backhurst (pers. comm.), editor of the Bulletin of the Society drew my
attention to Benson et al. (1970) who quote Broekhuysen (1967) and Mack-
worth-Praed & Grant (1952) who accept T. hypoleucos as a breeding species
while Moreau (1966) and Chapin (1939) rejected the breeding record of the
van Somerens and Meinertzhagen and thus Benson et al. (op. cit.) followed
suit and also rejected the record.
van Someren (1922, 1932) refers to this earlier breeding record but without
re-quoting the data on which this was founded and doubtless the above-men-
tioned writers have not accepted the bare statements of these papers. However
I would draw attention firstly to the 191 1 work, "Studies of Birdlife in
Uganda", where a photograph is reproduced, showing a bird on its nest in
Uganda. The photograph was taken by Dr. R. A. L. van Someren and he des-
cribes how this was achieved and further adds "breeding is during August and
September". He describes the eggs. This statement was quoted by Mackworth-
Praed & Grant (op. cit.). The volume of photographs and text is probably
not readily available and was perhaps overlooked.
Secondly van Someren (1936) writes of the Sandpiper "also resident in
East Africa". He describes the nestling in down and under habits comments
"is to a certain extent resident in both Kenya and Uganda and even breeds
here" ..." nest and young have been recorded in Uganda in June and a bird
in breeding condition shot in May" . . . "we have taken or observed these
Sandpipers throughout all months of the year on Lakes Nakuru and Naiv-
asha". Neither of these two papers are mentioned by Benson et. al (op. cit.).
The fact that T. hypoleucos has been photographed on its nest in Uganda,
the eggs and nestling described, etc., establishes the claim that it should now
be considered an East African breeding species. It is certainly resident
throughout the year in Kenya, as my own records show for my dam at
Karen, near Nairobi.
References :
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Stewart Irwin, M. P. 1970. Notes on the
birds of Zambia: Part V Arnoldia Vol. 4 No. 40.
Broekhuysen, G. 1967. Bird Migration in the southern part of the African continent.
Vogelwarte, 24: 6-16.
Chapin, J. P. 1939, 1954. The Birds of the Belgian Congo. Bull. Amer. Mus. Nat. Hist. 75,
75B.
Forbes-Watson, A. 1972. Birds naturalised in East Africa. Bull. E. A. Nat. Hist. Soc,
August.
Mackworth-Praed, C. W. & Grant, C. H. B. 1952. The birds oj eastern and north-eastern Africa.
I. London: Longmans.
Moreau, R. E. 1966. The bird faunas oj Africa and its islands. London: Academic Press,
van Someren, R. A. L. & van Someren, V. G. L. 191 1. Studies of Birdlife in Uganda. London:
John Bale, Sons and Danielson.
van Someren, V. G. L. 1922. Notes on the Birds of East Africa. Novitates Zoologicae Vol.
XXIX.
— 1932. Birds of Kenya and Uganda, being addenda and corrigenda to my previous paper
in Novitates Zoologicae. Novitates Zoologicae Vol. XXXVIII.
— 1936. The Birds of Kenya and Uganda. Jour. E. A. and U. Nat. Hist. Soc. Vol. XIII
Nos. 1 and 2. Vol. II Part 5.
40
Seasonal movements of the black cuckoo-shrikes
Campephaga phoenicea and C. flava, especially
in eastern Africa
by P. L. Br it ton
The Campephaga phoenicea superspecies of Hall & Moreau (1970) ranges widely
through Africa represented by three forms which are considered conspecific
by White (1962). Two of these, C. phoenicea (Latham) and C. flava Vieillot,
are apparently very tolerant ecologically, preferring woodland and savanna
habitats, but found also in forest. The third form, C.petiti Oustalet, is a true
forest bird confined to the more equable lower latitudes, and probably
resident. The plots for all three forms in Hall & Moreau (pp. cit.) indicate
eastern and western populations of each. Two of the forms are sympatric in
several areas, with considerable overlap o£ phoenicea and flava in the east. All
are more or less sympatric near the Equator in eastern Zaire and western
Kenya, and in the latter area the situation is further complicated by the
presence of the closely related, forest-dwelling C. quiscalina Finsch. Phoenicea
is a northern form ranging from about 14 N to 4 S in the west, 16 N to
1 ° S in the east ; while flava is a southern form ranging from about 5 ° S to
22 S in the west, 5 N to 34 S in the east.
There is considerable controversy and confusion surrounding the taxo-
nomic treatment of the two non-forest forms. The otherwise black males
usually have brightly coloured shoulder patches, most often red in phoenicea
and yellow in flava; but this aspect is not at all consistent for either form, not
even at one locality. In eastern Africa the vast majority of male flava are
entirely black, especially in central and southern Tanzania and southern
Uganda. About half of the males in several areas of Kenya and northern
Tanzania have yellow shoulders, but at Kabete, near Nairobi all eighteen
birds checked for this character by G. C. Backhurst (in litt.) were entirely
black. An analysis of any regular movements in these forms should help in
solving this complex problem; in particular whether areas of apparent
sympatry are occupied by only non-breeding birds.
The Common Black Cuckoo-Shrike C. flava was not included as a migrant
by Moreau (1966). Yet as early as 1936 Priest had remarked that it is a breed-
ing summer visitor to the Mashonaland plateau of Rhodesia; a view strongly
supported by Vernon (1968) who suggested that the movement is altitudinal.
Benson et al. (1970) showed that it is virtually absent from south-eastern
plateau country, especially Zambia, from June to September. During the
same period it is quite numerous below 3,500 ft.; a pattern which led the
authors to believe that the great majority on the plateaus leave for the winter,
moving generally eastwards into the low-lying valleys and Mozambique.
Most recently, Prigogine (1972) has shown conclusively that in Zaire, ex-
cluding Katanga, it is a seasonal visitor from May to October. Prigogine
considered it possible that some at least of these birds originate from Katanga
while others originate from East Africa.
In order to investigate the situation in eastern Africa, dated records have
been collected from resident observers and museum specimen labels, with
the kind co-operation of many, all of whom are acknowledged later. The 710
dated records from southern Ethiopia, Kenya, Tanzania and southern
Uganda are shown by months in Table 1. Monthly totals for this large area
41
TABLE 1
Distribution of Campephaga flava records in eastern Africa by months
J F M A M J Jl A S O N D Total
Tanzania south of 7°S,
west of 38°E, Songea
excluded
5
5
3
1
-
-
-
-
1
-
-
1
16
Songea District, S. Tanzania
12
3
2
1
2
2
-
-
2
12
10
16
62
Uganda ; Tanzania west of
33°E
1
2
1
1
2
5
11
9
2
-
1
1
36
S.W. Kenya, 0°-3° S,
34°-36° E
2
-
-
1
3
6
14
7
3
16
-
3
55
Central Tanzania, 4°-7°S,
33°-38°E
2
1
-
6
1
4
-
5
5
3
4
1
32
"Coast", south of 4°S,
east of 38°E
1
-
5
3
3
5
6
12
18
18
5
3
79
"Coast", north of 4°S,
east of 38°E
-
3
3
5
2
8
9
31
2
1
2
8
74
Kenya & N. Tanzania, north
of 4°S, 36°-38°E, Nairobi &
Kabete excluded
11
10
12
6
10
15
14
23
19
10
3
7
140
Kabete, near Nairobi
1
3
-
2
1
2
8
11
15
28
1
-
72
Nairobi area, Kabete
excluded
2
3
3
6
8
4
10
17
16
11
5
2
87
Kenya, 0°-2°N, 34°-36°E
3
5
1
1
-
6
13
2
8
12
3
3
57
Total north of 7°S
23
27
25
31
30
55
85
117
88
99
24
28
632
Total south of 7°S
17
8
5
2
2
2
-
-
3
12
10
17
78
Total 40 35 30 33 32 57 85 117 91 111 34 45 710
show that 70 per cent of records from north of 7 S fall in the months June
to October, whereas 78 per cent of records from south of 7 S fall outside
this period. Since the more or less coincident period from May to October
is the time of the invasion in much of Zaire (Prigogine op. cit.) it is probably
unreasonable to suppose that Zaire birds originate from anywhere but the
south. Furthermore, the absence from south-eastern plateau country outlined
above, documented in particular for Rhodesia and Zambia, more or less
coincides with this invasion.
In the eight months from October to May, Benson et al. (op. cit.) had 300
specimen records from plateaus above 3,500 ft., compared with only about
100 from the whole area considered, including the plateaux, for the four
months June to September. The fact that there were fewer data per month
in the latter period is probably partly the result of less collecting in low-
lying Mozambique than in either Rhodesia or Zambia; but the difference may
also be due to a departure of some of these plateau-breeding birds from the
whole area considered between June and September, not simply a movement
within the area. It is likely that a proportion of individuals move into low-
lying areas like Mozambique while others move to lower latitudes, into Zaire
and eastern Africa. The Katanga region of southern Zaire is geographically
intermediate which accounts for the more or less even distribution of records
through the year (Prigogine op. cit.). Katanga data probably refer to both
breeding birds and wintering birds, including passage migrants, and do not
necessarily imply that birds in this area are resident.
42
The movements of Cossypba natalensis hylophona Clancey described by
Britton (1971) are somewhat analogous since eastern Rhodesia plateau
breeders move eastwards into Mozambique from about April to August, at
which time birds breeding in southern and south-western Zambia, and
adjacent areas, move northwards into Katanga and Moxico (Angola). These
two breeding populations are quite disjoint so that it is comparatively easy
to assign disjoint wintering areas to them. This is not the case with C. flava
however, so that it is not possible to know which move to lower altitudes
and which move to lower latitudes in the cold weather (dry season).
Non-coastal Tanzania south of about 7 S may be considered a part of the
south-eastern plateau, receiving most of its rain from December to April
(Griffiths 1958). All but one of the records from this area, Songea excluded,
are for these months, when Lynes (1934 and specimen labels in B.M.[N.H.])
recorded considerable gonad activity in the majority of his specimens from
Iringa and the Njombe Highlands. The data for Songea District, lying south
of io° S, are those of I. H. Dillingham {in lift,) who was resident there from
March 1 9 5 4 to July 1956. The bulk of records (80 per cent) refer to the period
October to January, which are the months of egg-laying in adjacent Malawi
and Zambia (Benson et al. 1964). Dillingham had no records for July or
August, and returning from leave on 29th May 1955 he did not record this
species until 20th September.
Southern Uganda and Tanzania west of 33 E, mainly the north-west,
are peripheral to Prigogine's area, and evidently receive the same invasion
from the south, with 26 out of 35 records in the three months June to
August. Similarly south-western Kenya and adjacent Tanzania from the
Equator to 3 S, 34 to 3 6° E, where 46 out of 5 5 records are for the months
June to October (only three in September). In central Tanzania, 4 to 7 S,
33 to 3 8° E, there is evidence of passage birds, with 28 out of 32 records in
the months March to May and August to November, and none in July which
is the peak month to the north-west. In much of eastern Africa and Zaire the
situation is complicated by the presence of breeding birds, but breeding is
l unknown and probably rare in the foregoing areas.
In coastal Tanzania, south of 4 S, 33 to 3 8° E, an area which includes also
extreme south-eastern Kenya, the majority (48 out of 79) of records fall in
; the months August to October. There are however records for all months
' except February, and breeding is known from this area. Sclater & Moreau
(1933) mention a nest with naked nestlings at Amari on 26th October, and
Vaughan (1930) considers it probably resident on Zanzibar with a breeding
season about October. In fact all three December records refer to the Usam-
bara Mountains which include Amani, so that there is only one record for
the coastal lowlands in the months December to February. In the mainly low-
lying areas of eastern Kenya, Ethiopia and Somalia, east of 3 8° E, north of
4 S, 65 per cent of records are for the months of June to August, especially
August with 3 1 out of a total of 74 records. All months except January are
represented, with as many as eight in December. Almost half of the eastern
Tanzania records are for the months September and October compared with
only three out of 74 for these months further north in eastern Africa; a
situation suggesting a southern origin for many individuals, returning south
fairly early and hardly occurring north of 4 S after the peak month of August.
East of 3 8° E it hardly occurs north of z° S, and the population in south-
eastern Ethiopia at about 5 ° N is an anomaly mentioned in more detail later.
43
During 25 years as a resident of Korogwe in north-eastern Tanzania,
Archbold (1972) has recorded it only three times, in September, October and
November.
The remaining, mainly highland areas of Kenya and northern Tanzania
show the same June to October peak, but complicated by the presence of
breeding birds and possible altitudinal movements within these areas. J. S. S.
Beesley {in litt.) found three nests between 1,500 and 1,700 m a.s.l. in the
Arusha National Park, northern Tanzania, indicating egg-laying in December
January and April. Two eggs from Emali Hill, Sultan Hammud, Kenya col-
lected by V. G. L. van Someren on 30th March 1940 were from one of
several nests found on a visit to this area (G. R. Cunningham-van Someren
in lift.). The same writer informs me of a nest at Karen, near Nairobi in May.
A female with a large shell-less egg in its oviduct collected at Eldama Ravine
on 4th April, and reported by Jackson (1938), is the only further Kenya
breeding record. According to Urban & Brown (1971) it has been recorded
breeding in Ethiopia in April and possibly in February, the latter being a bird
with testes greatly enlarged collected at Yavello (B.M. [N.H.] collection).
All of the above breeding sites are at about 1,500 m (a little under 5,000 ft.)
or higher.
The fact that 62 per cent of the 3 5 1 records for this large area of Kenya and
northern Tanzania are for the four months July to October strongly suggests
that some at least are birds breeding in the southern tropics. No month lacks
records completely however due to the presence of a minority of breeding
birds. At Kabete, near Nairobi the observations of G. C. Backhurst {in litt)
and others over several years show a still clearer pattern with 62 out of 72
records for the months July to October. Surprisingly, 28 of these are for
October, very late if they are indeed southern breedej s. Some at least may be
birds breeding at higher elevations in Kenya and northern Tanzania from
about November to January, wandering to lower areas during the cold,
damp months. The pattern is clear at other sites also, for example at Naivasha
in the Rift Valley where all records are in July (three), August (four) and
October (three).
In view of the April to October pattern of occurrences in south-western
Zaire, documented by Prigogine {op. cit.), western birds probably move
mainly northwards into this and adjacent areas during the dry season.
Further south in Angola this is evidently a bird of plateau country breeding
in October (Traylor 1963). It is unlikely that there is any regular altitudinal
movement in Angola in view of the total lack of records from the coastal
lowlands. I have not attempted to investigate the movements of western
birds in any detail.
The Red-shouldered Cuckoo-Shrike C. pboenicea has been known to be
migratory in parts of West Africa for some years, but regular movements
have not been suspected elsewhere. Bannerman (1953) considered it partially
migratory in Gambia, Sierra Leonne and Ghana, and Elgood (1959) con-
sidered it the only undoubtedly migratory African passerine at Ibadan,
Nigeria. Elgood et a/, (in press) have little doubt that it is truly migratory in
western Nigeria, though not so in the east where, for example, it occurs
around Enugu throughout the year. In the west it breeds in the north during
the rains, retreating to the more equable lowland forest belt (for example at
Ibadan) from about November to April.
44
Table 2 shows dated records of C. phoenicea from throughout its range.
Sight records have been included for Ethiopia, Uganda and Kenya; elsewhere
TABLE 2
Distribution of C. phoenicea records by months
J F M A M J Jl A S N D Total
Nigeria (9) & west (26),
north of 7^ N -1638944---- 35
Nigeria (2) & west (10),
south of 7i°N 17__2------2 12
Cameroon 31-1221----- 10
Sudan & Zaire 553515222--1 31
Ethiopia & Somalia 9 6 7 6 10 10 3 4 1 1 8 19 84
Uganda & W. Kenya 888954534345 66
Total 26 28 24 24 28 30 15 13 7 4 12 27 238
specimens only. Data from Nigeria and further west are split into those
respectively north and south of 7^° N. In this area the northern limit of low-
land forest varies from about 6° N to 9 N, rendering 7-J N a somewhat
arbitrary line. Nevertheless a clear pattern emerges, with 34 out of 35
northern specimens between March and August, and 10 out of 12 southern
specimens between December and February (the two in May are from the
Prah River, southern Ghana). Bannerman {op. cit.) noted that it is seen only
during the northern winter at Kumasi in southern Ghana, while in Sierra
Leone it is more often seen from May to December. Sierra Leone specimens
in Table 2 were collected in February, March (three), Mat and June (five),
and all refer to the lowland forest belt, here extending to about 9 N. Some
West African birds are no doubt more or less resident or at most wanderers,
but the well-documented Nigerian pattern of a northward movement during
the wet summer (breeding) months of March or April to September or
October is widely applicable west of about 8° E.
Data from east of Nigeria show no clear pattern. The fact that only 21 per
cent of records from throughout its range refer to the five months July to
November invalidates any inference drawn from the comparative lack of
records east of Nigeria during this period. This is especially true since these
months follow the bulk of breeding in the northern tropics, so that if any-
thing numbers should be swollen by the presence of young birds in the popu-
lation. Such a lack represents either a lack of collecting during these months
or an overall movement into little-worked areas. No pattern was evident
when the records were accurately mapped, nor when areas like Ethiopia were
split into highland and lowland regions.
Despite the lack of an overall pattern east of 8° E it is likely that this
species wanders somewhat randomly over considerable distances, with a
minority performing regular movements comparable to those further west.
Ignoring the atypical western highlands of Ethiopia which extend north to
about 1 6° N, the most northerly specimens are from about 12 N in the
Sudan. Since they were taken by only three collectors these few specimens
represent a biased sample, referring to April, June (two), July (four),
August (five) and September. Despite the bias they indicate a northward
movement during the summer rains, when Lynes (1925) recorded it breeding
in July in the Darfur region of western Sudan. In contrast the unbiased
records of Rev. P. J. Hamel {in litt.) and other residents of the Kampala area
45
of southern Uganda, virtually on the Equator, are mainly (24 out of 29) for
the four months January to April, peaking in February (nine records). Some
at least of these records probably refer to non-breeding visitors from the
north, although it is known to breed in the south of its range also. Seth-Smith
(191 3) collected a clutch at Mpumu in southern Uganda on 3 1st March; van
Someren (19 16) collected a clutch at the Sezebwa River in southern Uganda
in April; and Tennent (1965) watched a pair building at Kakemaga Forest in
western Kenya in early June.
The important areas of apparent sympatry oi flava and phoenicea are south-
western Uganda, extreme western Kenya between Mt. Elgon and the Kavir-
ondo Gulf, southern Ethiopia, and southern Sudan (a single male flava col-
lected at Gilo). It has already been shown that southern Uganda flava are
off-season birds breeding further south. In Central Nyanza, western Kenya,
where I have been resident for over four years, I have recorded phoenicea only
six times, in January, June, July (three) and October. These and the few
other western Kenya records hardly overlap geographically with, flava which
occurs only to the north and east, and not to the north-west of this area,
there being no Uganda records of flava east of Kampala. In the area of
apparent sympatry (Central Nyanza, Kakamega, Kaimosi), 91 per cent of the
34 flava records fall in the months May to October when southern breeding
birds probably penetrate beyond the Equator; so that the vast majority of
flava may be considered non-breeders in the area of sympatry. The few
Ethiopian specimens of flava are an anomaly since the only other specimens
north of 2 N are from Kenya between z° and 3^-° N in March, June (three),
July (two), and a single specimen from southern Sudan in March. The
scattered Kenya birds are probably mainly southern breeders but the few
Ethiopian birds ate likely to be more or less resident in view of two breeding
records and the fact that they refer to the months February, June, July,
September and December. This isolated population occurs east of the Rift,
the male specimens available to me being from around 5 N. Only a single
Ethiopian specimen of phoenicea is from south of the restricted flava area, and
the vast majority of Ethiopian phoenicea are from the Rift and further north-
west. Immatures and females have been ignored in critical areas, being mainly
indeterminable.
Prigogine {op. cit.) has shown that the two closely allied forms, C. flava and
C. petiti, overlap geographically in Zaire at a time when neither is breeding.
In the areas of sympatry of phoenicea and flava the latter is a non-breeding
visitor except in south-eastern Ethiopia where the few phoenicea are probably
wanderers. The single Kenya nesting record of phoenicea is from an area where
petiti is common, but there is no competition between these forms since
petiti is found only in true forest. The phoenicea were building in an isolated
forest patch.
SUMMARY
Account has been taken of 710 dated records of C am pephaga flava from
eastern Africa, and of 238 dated records of C. phoenicea from throughout its
range, with a view to establishing the nature of any movements.
C. flava breeding in southern Africa north to about 7 S in the east, 9 or
io° S in the west, are mainly not resident, leaving plateau country during the
dry months of May to September or October. Some move to lower altitudes
at similar latitudes, especially into low-lying Mozambique, while others move
north to lower latitudes. The latter account for the majority of records in
46
Zaire (excluding Katanga), Tanzania north of 7 S, southern Uganda and
Kenya, exceptionally north to 3 N but mainly north to about the Equator.
There are locally breeding, morphologically indistinguishable birds in high-
land areas of eastern Zaire, northern Tanzania and Kenya which probably
move altitudinally to some extent. A localized population in south-eastern
Ethiopia is an anomaly and probably mainly resident.
C. phoenicea is migratory in west Africa east to about 8° E, moving north
from the lowland forest belt to breed during the rains from March or April to
September or October. Further east movements are not clear-cut except at
the northern and southern extremities of its range, in the Sudan and southern
Uganda respectively, where the pattern of records follows that established
for West Africa. Elsewhere movements are probably in the nature of random
wanderings. It breeds locally throughout its range.
In the areas of apparent sympatry of these two non-forest forms in eastern
Africa, flava is a non-breeding visitor except in south-eastern Ethiopia where
the few phoenicea are probably wanderers. It is unlikely that they ever
interbreed.
ACKNOWLEDGEMENTS
I am very grateful to the following who supplied personal records : Miss
P. M. Allen, Dr. J. S. Ash, G. C. Backhurst, J. S. S. Beesley, A. Beverly,
Miss E. J. Blencowe, Mrs. S. Brashaw, G. R. Cunningham-van Someren,
I. H. Dillingham, Dr. P. Duffus, Mrs. F. Edwards, L. A. Haldane, Rev.
P. J. Hamel, W. G. Harvey, Dr. W. Leuthold, C. F. Mann, P. J. Oliver,
J. F. Reynolds, V. R. Simpson, Dr. N. J. Skinner, C. Smeenk, Dr. E. K.
Urban. The following kindly supplied data on the specimens in the institu-
tions shown in parenthesis after their names: Prof J. Berlioz (Museum
National d'Histoire Naturelle, Paris), G. M. Bond (Snithsomian Inst.), Dr.
F. B. Gill (Academy of Natural Sciences, Philadelphia), Dr. J. van der Land
(Leiden Museum), K. C. Parkes (Carnegie Museum, Pittsburg), Dr. R. B.
Payne (Museum of Zoology, University of Michigan), Dr. R. A. Paynter
(Museum of Comp. Zoology, Harvard), Dr. F. Salomonsen (Universitetets
Zoologiske Museum, Copenhagen), K. E. Stager (Los Angeles County
Museum), Major M. A. Traylor (Field Museum, Chicago), Dr. H. E. Wolters
(Zoologisches Museum Alexander Koenig, Bonn.) Rev. P. J. Hamel kindly
sent records from the monthly duplicated newsletter of Uganda biid records,
1 969-1 972; and Dr. C. H. Fry kindly sent details of C phoenicea from Elgood
et al. (in press). I have used the unpublished notes of A. F. Morrison lodged
in the National Museum Library, Nairobi. The Directors of the British
Museum (Natural History) and the National Museum, Nairobi kindly allowed
me access to their collections.
References :
If published data were also supplied in a list of specimens the reference is not given. Those
marked with an asterisk are data sources not mentioned in the text.
Archbold, M. E. 1972. Exceptional bird activity at Korogwe, Tanzania. Bull. E.A.N.H.S.,
1972: 45-47.
*Bannerman, D. A. 19 10. On a collection of birds made by Mr. A. B. Percival in British
East Africa. Ibis 4 (9): 676-710.
— 1953. The Birds of West and Equatorial Africa. Edinburgh & London: Oliver & Boyd.
Benson, C. W., Brooke, R. K. & Vernon, C. J. 1964. Bird breeding data for the Rhodesias
and Nyasaland. Occ. Pap. natn. Mus. Sth. Rhod. 27B: 30-105.
47
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. S. 1970. Notes on the birds
of Zambia: part V. Arnoldia {Rhodesia) 4 (40).
Britton, P. L. 1971. On the apparent movements of Cossypha natalensis. Bull. Brit. Orn. CI.
91: 137-144.
Elgood, J. H. 1959. Bird migration at Ibadan, Nigeria. Ostrich, Suppl. 3: 306-316.
Elgood, J. H., Fry, C. H. & Dowsett, R. J. in press. African migrants in Nigeria. Ibis.
*Friedmann, H. 1937. Birds collected by the Childs Frick Expedition to Ethiopia and Kenya
Colony. Part 2, Passeres. Bull. U.S. Nat. Mus. 153.
*Friedmann, H. & Loveridge, A. 1937. Notes on the ornithology of tropical East Africa.
Bull. Mus. Comp. Zool. Harvard 81 (1).
Granvik, H. 1923. Contributions to the knowledge of East African ornithology. Birds
collected by the Swedish Mount Elgon Expedition, 1920./. Orn. 71 : 1-280.
Griffiths, J. F. 1958. The climatic zones of East Africa. E. Afr. Agric.Journ., 1958: 179-185.
Hall, B. P. & Moreau, R. E. 1970. An atlas of speciation in African passerine birds. London:
British Museum (Nat. Hist.).
Jackson, F. J. 1938. The Birds of Kenya Colony and the Uganda Protectorate. London:
Gurney & Jackson.
Lynes, H. 1925. On the birds of north and central Darfur (part). Ibis 12 (1): 71-131.
— 1934. Contribution to the ornithology of southern Tanganyika Territory. Birds of the
Ubena-Uhehe highlands and Iringa uplands.
/. Orn. 82: 1-147.
Moreau, R. E. 1966. The bird faunas of Africa and its islands. London: Academic Press.
*Norris, E. 1970. Letter. Newsletter E.A.N.H.S., 1970: 22.
Priest, CD. 1936. The birds of Southern Rhodesia. 4. London: William Clowes.
Prigogine, A. The seasonal migrations of the Common Black Cuckoo-Shrike Campephaga
f'ava Viellot. Bull. Brit. Orn. CI. 92: 83-90.
Sclater, W. L. & Moreau, R. E. 1933. Taxonomic and field notes on some birds of north-
eastern Tanganyika Territory (part). Ibis (13) 3: 187-219.
Tennent, J. R. M. 1965. Notes on the birds of Kakamega Forest. //. E. Afr. nat. Hist.
Soc. 25 (2): 95-100.
Traylor, M. A. 1963. Checklist of Angolan birds. Publ. Cult. Comp. Cia. Diamant. Angola 61.
*Tweedy, C. J. 1966. A list of birds seen in southern Tanzania. Tang. Notes <& Rec. 25(3):
179-188.
Urban, E. K. & Brown, L. H. 1971. A checklist of the birds of Ethiopia. Addis Ababa:
Haile Selassie I Un. Press,
van Someren, V. G. L. 1916. A list of birds collected in Uganda and British East Africa,
with notes on their nesting and other habits. Ibis. (10) 4: 193-252, 373-472.
* — 1 921. On a collection of birds from Turkanaland. //. E. Afr. <& Uganda nat. Hist. Soc.
16:3-38.
* — 1928. Notes on the birds of Jubaland and the Northern Frontier, Kenya. //. E. Afr. &
Uganda nat. Hist. Soc. 35 : 25-70.
Vaughan, J. H. The birds of Zanzibar and Pemba (part). Ibis (12) 6: 1-48.
Vernon, C. J. 1968. A year's census of Manardellas, Rhodesia. Ostrich 39: 12-24.
White, C. M. N. 1962. A revised checklist of African shrikes, etc., Lusaka: Govt. Printer.
* Williams, J. G. 1947. Notes on the monthly rambles of the E.A.N.H.S. Nat. E. Afr. 3 : 22.
48
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Published by the BRITISH ORNITHOLOGISTS' CLUB and printed by
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-•»• n
NAT
HIST ^
2 6JUNI973 1
Bulletin of the
British Ornithologists' Club
Edited by
C. W. BENSON
Volume 93 No. 2
June 1973
Committee
Sir Hugh Elliott, Bt., o.b.e. {Chairman)
J. H. Elgood {Vice-Chairman)
C. W. Benson, o.b.e. {Editor)
R. E. F. Peal {Hon. Secretary)
P. Tate {Hon. Treasurer)
J. H. R. Boswall
Dr. P. J. K. Burton
P. Hogg
C. J. Mead
to
tew
Bulletin of the
BRITISH ORNITHOLOGISTS' CLUB
Vol. 93 No. 2 Published: 20 June, 1973
The six hundred and eighty-first meeting of the Club was held at the Cafe
Royal, 68 Regent Street, London, W.i, on Tuesday, 20th March, 1973 at
7P- m \
Chairman: Sir Hugh Elliott, Bt., o.b.e.; present 26 members and 22 guests.
The speaker was Mr. Guy Mountfort, o.b.e., who gave an illustrated
address on his recent visit to India, Pakistan and Nepal.
The six hundred and eighty-second meeting of the Club was held at the
Cafe Royal, 68 Regent Street, London, W.i, on Tuesday, 8th May, 1973 at
7 p.m.
Chairman: Sir Hugh Elliott, Bt., o.b.e.; present 16 members and seven
guests.
The speaker was Mr. Tom Harrisson, d.s.o., o.b.e., who opened with
some amusing recollections of meetings of the Club at the time when he had
last spoken, 40 years previously. He then gave an illustrated address on the
birds and bird cult of Easter Island and a fuller report on this will be pub-
lished in a future number of the Bulletin.
Annual General Meeting
The eighty-first Annual General Meeting of the British Ornithologists'
Club was held at the Cafe Royal, 68 Regent Street, London, W.i, on Tues-
day, 8th May 1973 at 6 p.m. with Sir Hugh Elliott, Bt., o.b.e., in the Chair.
Eight members were present.
The Minutes of the 1972 Annual General Meeting [Bull. Brit. Orn. CI.
92 : 69-70) were approved and signed.
After a brief discussion on the Report of the Committee for 1972 {Bull.
Brit. Orn. CI. 93 : 1-2), Mr. C. J. Mead proposed and Mr. J. H. Elgood
seconded the adoption of the Report of the Committee for 1972 and this was
carried unanimously.
Presenting the Accounts for 1972 {Bull. Brit. Orn. CI. 93: 50-51) the
Hon. Treasurer stated that 1972 was notable for the unusually large receipts
from sales of back-numbers of the Bulletin. It was extremely unlikely that they
could be maintained at about £400 a year and in addition the printers had
just announced a 10% increase in theit charges for printing, which would
cost the Club about £90 in the current year and some £120 in a full year.
The charge for subscribers to the Bulletin had been raised in 1972 from £3 to
£3. 50, to take effect from 1st January 1973, which would bring in about £50
more, but an increase in membership was essential. The Hon. Secretary
stated that a leaflet, which it was hoped would produce more new members,
would be distributed in the July Ibis. Mrs. J. D. Bradley proposed and Mr.
J. H. Elgood seconded that the Accounts for 1972 be adopted and this was
carried unanimously.
There being no nominations additional to those of the Committee, the
following elections were made: —
Hon. Secretary: Mr. R. E. F. Peal (re-elected)
[continued page 52]
49
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Hon. Treasurer: Mr. P. Tate (re-elected)
Committee: Mr. J. H. R. Bos wall
The Editor stated that the publication of the Bulletin quarterly had pre-
sented no great difficulties. There were ample papers of a high standard being
offered but the average time from receipt to publication was only four
months.
The meeting closed at 6.25 p.m.
A recent immigrant to Ontong Java atoll,
Solomon Islands
by T. P. Bayliss-Smith
Received 2nd April, 1975
The work of MacArthur & Wilson (1967) is founded on the notion that on an
island there exists a tendency towards an equilibrium condition in its biota.
An equilibrial species number is reached at the intersection point between
the curve of rate of immigration of new species not already on the island, and
the curve of extinction of species from the island. The general idea of a
balance of immigration by extinction was originally suggested to these
authors by the impoverished bird faunas of the islands of central and eastern
Polynesia (MacArthur & Wilson 1963). The theory has received empirical
support from some remarkable experiments by Simberloff & Wilson
(1969—70) on the recolonization of depopulated mangrove islands by insects,
and from the work of Diamond (1969, 1971) on the turnover rates of the bird
populations of Californian and New Guinean islands. One disadvantage of
these studies is the cross-sectional character of the field data. The island
populations are surveyed at intervals in time, but little is known about what
happens in between surveys. The actual processes of immigration and
extinction must usually be deduced from considerations of island size and
distance from the source area.
It is of interest, therefore, to record an actual case of the colonization of an
island by a new bird species. The birds of Ontong Java atoll, Solomon
Islands, were observed over a one-year period from June 1970 to May 1971.
Eight species of land birds were recorded of which seven were breeding
residents and one, Eudynamis taitensis, was a seasonal migrant (Bayliss-Smith
1972). In July 1972 the atoll was revisited. A new species, Eos cardinalis, the
Cardinal Lory, was found to be present. A noisy flock of about twelve of
these long-tailed scarlet parrots was seen on several occasions on Luangiua
islet. Reliable native informants stated that these unmistakable newcomers
were present on all the islets between Keila and Henguakaha on the southern
and eastern sides of the atoll. They have been named "kesivi", and were first
seen after a period of south-westerly gales at the beginning of June 1972.
These winds were generated by Cyclone Ida, a small hurricane which formed
on May 31st over the ocean east of Bougainville and moved during the
following day southwards and eastwards across Choiseul and Santa Isabel,
causing much local destruction (British Solomon Islands 1972). Eos cardinalis
is exceedingly numerous throughout the Solomon Islands from Nissan to
Ugi, and "large flocks . . . are often seen flying from one island to the next"
(Mayr 1945 : 230). It must be assumed that one such flock was blown out to
5 2
sea by the hurricane and found refuge on Ontong Java, which lies 250 km
north-east of Choiseul and Santa Isabel.
Four months after its first arrival Eos appeared to be flourishing in its new
atoll environment. Although there were no reports of nesting, there seems
to be no reason why a breeding population should not become established.
The vegetation of the islets where it has settled is mainly coconut woodland,
a habitat very similar to the coastal coconut plantations of the Solomon
Islands where Eos is so numerous. There are already complaints by local
people of damage caused by the birds to the flowers of coconut palms. The
ecology of the species involved is not well known, but it is probable that
Eos will compete both for food and nest sites with the two Ontong Java
pigeons Ducula pacifica and Caloenas nicobarka, and possibly also with the
Atoll Starling Aplonis feadensis. It will be interesting to see in the future
whether the atoll can accommodate the newcomer without the displacement
of one of the existing residents, or whether a total of eight land birds is in
fact the equilibrium species number for an atoll of this size.
References :
British Solomon Islands : 1972. The fury of Cyclone Ida. BSI News Sheet 1972 (11) ,1-15
June. Honiara: Government Printer.
Bayliss-Smith, T. P. 1972. The birds of Ontong Java and Sikaiana, Solomon Islands. Bull.
Brit. Orn. CI. 92: 1-10.
Diamond, J. M. 1969. Avifaunal equilibria and species turnover rates on the Channel
Islands of California. Nat. Acad. Sci., Proc. 64: 57-63.
Diamond, J. M. 1971. Comparison of faunal equilibrium turnover rates on a tropical island
and a temperate island. Nat. Acad. Sci., Proc. 68: 2742-2745.
MacArthur, R. H. & Wilson, E. O. 1963. An equilibrium theory of insular biogeography.
Evolution 17: 373-387.
MacArthur, R. H. & Wilson, E. O. 1967. The Theory of Island Biogeography. Princeton, New
Jersey: Univ. Press.
Mayr, E. 1945. Birds of the Southwest Pacific. New York: Macmillan.
Simberloff, D. S. & Wilson, E. O. 1969-70. Experimental zoogeography of islands. 1. The
colonization of empty islands. 2. A two-year record of colonization. Ecology 50:
278-296, and 51: 934-937.
The correct spelling of Scopoli's specific name for the
Maylaysian Crested Wood Partridge (Rollulus)
by John Farrand^ Jr. <& Storrs L. Olson
Received 16th February, iyjj
For more than a century the specific name of the Malaysian Crested Wood
Partridge has been subject to persistent mis-spelling. Some authors have
listed the species as Rollulus rouloul (Scopoli) but most, inserting a second r,
have cited it as Rol/us roulroul, still under the authorship of Scopoli. Basing
his description on "Le Rouloul de Malacca" of Sonnerat (1782: 174, 286,
pi. 100), Scopoli (1786: 93) named the bird Phasianus {Rouloul). Owing
perhaps to the rarity of Scopoli's publication, his name was subsequently
overlooked, generally in favour of Columba cristata Gmelin (1788, vol. 1,
pt. 2 : 774). The vernacular name "Rouloul" did, however, serve as the basis
for the genus Rollulus Bonnaterre (1791, vol. 1 : xciii).
Scopoli's name was resurrected by G. R. Gray (1844: 42; 1846, vol.
3: 507; 1870: 269) who unfortunately mis-spelled it roulroul. Since the
53
publication of these works by Gray, the spelling roulroul has appeared
frequently in the literature, and with the use of this spelling by Ogilvie-
Grant (1893 : 225), who evidently followed Gray, and by Peters (1934: 104),
the erroneous spelling has all but supplanted that appearing in Scopoli's
original description. Notable exceptions have been, among others, the
several papers on the birds of Borneo by R. B. Sharpe (1876: 51; 1879: 270;
1 881 : 800; 1890: 140) and by Salvadori (1874: 308), in which the bird is
listed as Kollulus rouloul. Baker employed both spellings in the second edition
of the Fauna of British India, calling the species Kollulus roulroul in Volume 5
(1928: 368) and Kollulus rouloul in Volume 7 (1930: 459). More recently, the
original spelling of Scopoli has been used by Smythies (i960: 168; 1968:
172). Most other works published within the last fifty years have listed the
species as Kollulus roulroul.
We are not aware of any published, formal emendation of Scopoli's name.
Indeed, such an emendation would not be justified, since there was no error
on Scopoli's part. He based the name on "Le Rouloul" of Sonnerat, who
clearly intended this spelling, as it reappears in the second edition of his
Voyage (1806: 300). It is evident therefore that Scopoli's original spelling
should be used and that the Malaysian Crested Wood Partridge should be
referred to as Kollulus rouloul (Scopoli). This spelling may be adopted without
application to the International Commission on Zoological Nomenclature,
since it has been used within the last fifty years, and therefore is not a nomen
oblitum as defined in Article 23b of the 1961 and 1964 International Code of
Zoological Nomenclature (see also Corliss 1972).
We would like to thank S. Dillon Ripley and George E. Watson for
helpful suggestions during the preparation of this manuscript, and Jack F.
Marquardt for making it possible for us to examine an original copy of
Scopoli's Deliciae in the Smithsonian Institution libraries.
References :
Baker, E. C. S. 1922-1930. The Fauna of British India. Birds. 2nd ed. 8 vols. London: Taylor
& Francis.
Bonnaterre, J . P. 1 790- 1823. Tableau Encyclopedique et Methodique des Trois Regnes de la Nature.
Ornithologie . 4 vols. Paris. (Pages lxxxi to scvii and 1 to 192 of Volume 1 were published
in 1791.)
Corliss, J. O. 1972. Priority and stability in zoological nomenclature: resolution of the
problem of Article 23b at the Monaco Congress. Science 178: 1120.
Gmelin, J. F. 1788-1793. Systema Naturae. 13th ed. 3 vols. Leipzig.
Gray, G. R. 1844. List of the Specimens of Birds in the Collection of the British Museum, Part III.
London: British Museum.
— 1 844-1 849. The Genera of Birds. 3 vols. London : Longmans, Brown, Green & Longmans.
(Pages 504 to 508 of Volume 3 were published in January 1846.)
— 1870. Hand-list of Genera and Species of Birds, Part II. London : British Museum.
Ogilvie-Grant, W. R. 1893. Catalogue of the Game Birds in the Collection of the British Museum-
London (Cat. Birds Brit. Mus., Vol. XXII).
Peters, J. L. 1934. Check-list of the Birds of the World. Vol. 2. Cambridge, Mass.: Harvard
University Press.
Salvadori, T. 1874. Catalogo sistematico degli uccelli di Borneo. Ann. Mus. Civ. Stor. Nat.
Genova 5 : 1-429.
Scopoli, G. A. 1786. Deliciae Florae et Faunae Insubricae. Pars II. Pa via.
Sharpe, R. B. 1876. Contributions to the ornithology of Borneo. Part 1. Ibis (3)6: 29-52.
— 1879. Contributions to the ornithology of Borneo. Part 4. Ibis (4) 3 : 233-272.
— 1 88 1. On the birds of Sandakan, north-east Borneo. Proc. Zool. Soc. London: 790-800.
— 1890. On the ornithology of northern Borneo. Part 6. Ibis (6) 2: 133-149.
Smythies, B. E. i960. The Birds of Borneo. Edinburgh & London: Oliver & Boyd.
— 1968. The Birds of Borneo, 2nd ed. Edinburgh & London: Oliver & Boyd.
Sonnerat, P. 1782. Voyage aux Indes Orientales et a la Chine. 2 vols. Paris.
— 1 806. Voyage aux Indes Orientales et a la Chine. Nouv. ed. 4 vols. Paris.
54
Notes on a Lanner with malformed bill, and on
hornbills feeding on oil palm fruits
by W. R.J. Dean
Received 21st March, 1975
In August 1970 an immature Lanner Falco biarmicus was found dead on the
national road 16 km north-east of Christiana, Transvaal, South Africa. The
bird was made into a study skin and is now in the collection of the Alexander
McGregor Memorial Museum, Kimberley (specimen no. NMK/B/1180). It
has the upper and lower mandibles malformed as shown in the Plate. The
upper mandible curves down sharply and is twisted laterally to the right
crossing the lower mandible which has projected beyond the upper by
5 mm. The lower mandible is also twisted laterally. On dissection the bird
Immature Lanner with malformed bill.
proved to be in fair condition with a small amount of fat present. The
severely abraded condition of the rectrices and primaries indicated that it had
been obtaining food by scavenging along roads.
Brooke & Jeffery (1972, Bull. Brit. Orn. CI. 92: 19) list several species of
birds, including three hornbills, taking nuts of the oil palm Elaets guineensis.
In the Quicama National Park, Luanda, at ca. 9 20'S., 1 3 46'E., the Crowned
Hornbill Tockus alboterminatus was recorded on seven occasions between 26
July and 13 August 1972 feeding on oil palm nuts, and the Yellow-billed
Hornbill T. flavirostris as twice taking oil palm nuts at the same time and
place. The stomach contents of a female Laughing Hornbill Bycanistes sharpii,
collected on 16 August 1972 in evergreen forest about 20 km north of
Cabinda, consisted of oil palm nut fibre and oil. I am grateful to the Peabody
Museum of Natural History, Yale University for the opportunity to make
these hornbill notes.
55
Diomedea cauta in South African waters
by C. M. N. White
Received 14th March, 197}
This albatross has long been recorded from the coastal waters of South
Africa in the form D. c. salvini which breeds at Snares Island and Bounty
Island, south of New Zealand. It now appears that the nominate D. c. cauta
which breeds on islets off Tasmania, and at Auckland and Disappointment
Islands, also occurs. My attention was drawn to this fact by the reference in
Checklist Committee, Orn. Soc. N. Z. (1970).
Dr. F. C. Kinsky of the Dominion Museum, Wellington, New Zealand
has kindly given me further details. Three birds are recorded, all banded at
sea off Cape Campbell, north-east corner of the South Island of New Zea-
land:
(a) No. 19289. Banded 4.3.1958: recovered off Port Nolloth, 3.5.1959.
(b) No. 19354. Banded 9.1. 1957: recovered off Danger Point, 30.8.1962.
(c) No. 16117. Banded 24.1. 1957: recovered off Jeffrey's Bay, 29.9.1968.
According to Dr. Kinsky the first two recoveries were published in the
10th and 13th Annual Reports of the New Zealand Bird Banding Scheme,
published by the Dominion Museum in i960 and 1963 respectively. The
third was mentioned by Robertson (1972: 71). Dr. Kinsky, to whom I am
much indebted for this information, comments that these birds were banded
during the New Zealand breeding season, and were probably sub-adult
non-breeding birds. Most of the nominate form are thought to remain in
New Zealand waters throughout the year.
Differential diagnoses of the forms can be found in Falla et al. (1966: 34)
and Serventy et al. (1971 : 70-71). This note is published in case students of
African sea-birds overlook these records, which provide an additional form
to the African marine bird fauna.
References :
Checklist Committee, Orn. Soc. of New Zealand, Inc. 1970. Annotated checklist of the birds
of New Zealand. Wellington: A. H. & A. W. Read.
Falla, R. A., Sibson, R. B. & Turbott, E. G. 1966. A field guide to the birds of "New Zealand.
London: Collins.
Robertson, C. J. R. 1972. Preliminary report on bird banding in New Zealand 1964-71.
Notornis 19(1): 61-73.
Serventy, D. L., Serventy, D. & Warham, J. 1971. The handbook of Australian sea-birds.
Sydney, Melbourne, Wellington, Auckland: A. H. & A. W. Read.
Winter bird populations of Golfo San Jose, Argentina
by J. R.JehlJr., M. A. E. Rumboll & /. P. Winter
Received 6th February, 1973
INTRODUCTION
Five hundred miles south of Buenos Aires, Argentina's Valdes Peninsula
juts into the Atlantic Ocean. Its irregular dumbell-like shape is caused by two
large bodies of water, Golfo Nuevo to the south and the smaller Golfo San
Jose on the north, which cut deeply into the peninsula and nearly separate it
from the mainland. The peninsula and its adjacent waters are attractive to a
variety of sea birds and marine mammals. Although there are nesting colonies
of cormorants, gulls, and terns there, the primary attractions are the large
herds of elephant seals and sea lions.
56
In the austral winters of 197 1 and 1972, the R/V Hero, a research vessel of
the U.S. National Science Foundation, supported biological field studies
along the coast of Argentina. During 22-23 J une an d 1-7 July 1971, and
4-19 August 1972, the Hero was engaged in bird and mammal research in
Golfo San Jose. This gulf is approximately 12 miles wide and 25 miles long.
Its entrance is narrow, and the swift currents and shoal waters there create
feeding conditions that attract many species of pelagic birds. In mid-gulf
depths reach 3 5-40 fathoms, but the waters within a mile or so of shore are
extremely shallow, and at low tide extensive mudflats a half-mile wide are
exposed in some areas. The perimeter of the gulf is dominated largely by
sandy beaches, some of which are thickly covered with layers of scallop
shells. Particularly along the southeast shore, steep, poorly-consolidated
sandstone cliffs drop directly into the water. In other areas, but mainly along
the north and south shores, there are scattered, dark-coloured mudstone
outcrops, rome of which are covered with mussel beds. The richest concentra-
tions of birds are usually encountered along the south shore of the gulf,
especially in the vicinity of Isla de los Pajaros. This tiny island provides
VALDEZ PENINSULA
Figure 1. Golfo San Jose on the Valdes Peninsula, Argentina: a, San Roman; b, Punta
Conos; c, Isla de los Pajaros. Cross-hatched areas along shore were censused in 1972.
Insert shows the location of the Valdes Peninsula.
undisturbed resting and feeding grounds for many shorebirds and water-
fowl, and in summer several species of aquatic birds nest there.
In 1 97 1, Jehl made preliminary censuses of birds along the entire south
shore and parts of the western shore; approximately 30 miles of coast were
surveyed casually. In 1972, we were able to make detailed censuses along 58
miles of the bay's approximately 80 miles of coastline. The areas censused are
shown in Figure 1. Censuses were made either by walking along the beach or
by observing from a small air boat that could operate within a few yards of
shore. By taking into account the types of habitat in areas we were unable to
visit, we could, through extrapolation, estimate populations for the entire
coast of the gulf. Numbers observed in areas censused more than once
showed little variation from day to day, indicating (1) a relatively stable
population at this season and (2) reasonable censusing techniques. We
believe that our estimates are accurate to within 10 per cent.
57
Although there are annotated lists of species for a few areas (e.g., Punta
Tombo: Boswell & Prytherch 1972; Puerto Deseado: Zapata 1967) ours are
apparently the only quantitative data on population sizes of shore-inhabiting
birds for any area of the Argentine coast at any season of the year.
TABLE 1
Winter bird populations of Golfo San Jose,
Argentina
1972
1971
1972
Estimate
Species
Estimate
Count
for entire gulf
Spheniscus magellanicus
100
250
400 +
Podiceps rolland
20
1
1
Podiceps occipitalis
5
43
50
Podiceps major
10
42
50
Diomedea melanophris
30
150
150
Macronectes giganteus
10
10
30
Fulmarus glacialoides
—
1
1
Daption capense
2
6
20
Pachyptila sp.
40
2,000
2,000
Procellaria aequinoctialis
2
60
80
Puffinus griseus
500
2,000
2,000
Puffinus gravis
10
—
—
Oceanites oceanicus
5
—
—
Phalacrocorax olivaceus
100
143
180
Phalacrocorax magellanicus
—
84
84
Phalacrocorax atriceps
50
53
70
Phalacrocorax albiventer
1,000 +
613
850
Casmerodius albus
2
4
4
Nycticorax nycticorax
50
68
75
Phoenicopterus chilensis
300
214
214
Chloephaga picta
3
40
40
Tachyeres patachonicus
8
23
25
Lophonetta specularoides
70
233
280
Anas flavirostris
6
6
6
Anas georgica
—
46
46
Falco peregrinus
—
1
1
Falco femoralis
1
—
—
Haematopus palliatus
500
492
600
Haematopus leucopodus
100
i75
175
Haematopus ater
8
22
22
Vanellus chilensis
—
1
1
Charadrius falklandicus
200 +
1,560
1,800
Zonibyx modestus
1
—
—
Pluvianellus socialis
30
10
10
Calidris canutus
1
36
36
Calidris bairdii
4
16
20
Calidris fuscicollis
50
7
10
Calidris alba
—
—
50
Thinocorus rumicivorus
—
50
50
Chionis alba
100
162
180
Catharacta skua
3
6
6
Leucophaeus scoresbii
4
2
2
Icarus dominicanus
1,000 +
4,300
4,800
Icarus maculipennis
50
466
500
Sterna hirundinacea
150
160
200
Sterna trudeaui
12
12
Thalasseus maximus
1
4
4
In each year, the Hero made several transects of the open gulf, and many
extended cruises along its shores. Estimates of pelagic species were made on
these occasions, but abundance varied markedly. For these species, the
number presented in Table 1 represents the maximum concentrations ob-
served in the census period.
For most species there were no important annual differences in abun-
dances, at least in those areas that received comparable study. It would
appear, therefore, that winter bird populations utilizing this area are largely
stable during the months of June, July, and August.
Surface water temperatures averaged io° C. in 1971 and 9 C. in 1972.
This research was supported by a grant from the National Science Founda-
tion (NSF-6V-32739).
WINTER BIRD POPULATIONS
Penguins: An estimated 400 Magellanic Penguins Spheniscus magellanicus , of
which approximately 10 per cent were immatures, wintered in Golfo San
Jose in 1972. They were widely scattered through the gulf, but concentra-
tions could usually be found near the mouth and along the southeast shore.
The largest concentration, 210 birds on 12 August, was observed in mid-gulf
in association with flocks of Sooty Shearwaters Puffinus griseus, prions, and
other pelagic birds. Two penguins collected had fed on a small fish Nothotenia
guntheri, and others that could not be identified. The prevailing westerly
winds cause birds dying in the bay to accumulate along the east shore. The
thick pelts of penguins resist decay, and one can find the mummified remains
of one or more every 30 metres along this sandy beach.
Grebes: Grebes were generally uncommon and tended to be concentrated
in small flocks near the south shore. The largest flock of Great Grebes
Podiceps major in 1972 was 25, and of Silvery Grebes P. occipitalis 15. These
concentrations were similar to those of 1971. Only one White-tufted Grebe
P. rolland was seen in 1972, as compared to a single flock of 20 in 1971.
Seabirds: Seabird numbers in Golfo San Jose fluctuated daily. Large
concentrations were usually present near the mouth of the gulf, but the
occurrence of concentrations elsewhere was much more variable. In most
cases we were unable to associate environmental factors with the presence of
particular species. However, on days of moderate to strong breezes large
numbers of Sooty Shearwaters appeared in the southeastern corner of the
gulf. It was not unusual to see 1,500 to 2,000 birds in that area, mostly
feeding on small fish a mile or so from the shore. On several occasions flocks
of several hundred to 2,000 fed at the surfline in association with Brown-
hooded Gulls Icarus maculipennis.
In general, the numbers and distribution patterns of seabirds showed
little variation between years. The apparent increased abundance in 1972
(Table I) largely reflects a huge concentration in mid-gulf on 12 August
which included 130 Black-browed Albatrosses Diomedea melanophris, 2,000
prions Pachyptila spp., 60 Shoemakers Procellaria aequinoctialis, 800 Sooty
Shearwaters, 130 Magellanic Penguins, 650 Kelp Gulls Larus dominicanus,
and 30 South American Terns Sterna hirundinacea. The 1971 data more closely
approximate the average daily numbers that can be expected.
The absence of Greater Shearwaters Puffinus gravis and Wilson's Petrels
Oceanites oceanicus in 1972, and the presence of a Southern Fulmar Fulmarus
glacialoides may be attributable to the later date and colder waters in that year.
We were unable to collect any of the prions for positive identification; a
single Pachyptila belcheri was found freshly dead on the beach.
Cormorants: Four species of cormorants winter, and one breeds in Golfo
San Jose. A colony of Olivaceous Cormorants Phalacrocorax olivaceus nests on
Isla de los Pajaros (see Ragonese & Piccinini 1972; Figs. 14, 16, 21, 23). In
winter the species is usually encountered feeding in very shallow water, in
areas where the bottom is sandy, mainly along the south shore.
59
The Rock Shag's P. magellanicus distribution is localized, and for this
reason the species was not encountered in 1971. Roosts are located on the
east and north shore, where steep cliffs plunge directly into the water. The
major roost, of about 50 birds, is at Punta Conos (Fig. 2); smaller roosts occur
Figure 2. Rock Shags Phalacrocorax magellanicus at roost at Punta Conos, Golfo San Jose.
near the mouth of the gulf and at Isla de los Pajaros. Rock Shags feed near
shore at depths of 10-20 metres. Several birds collected had fed on small fish
and on a small octopus Octopus tehuelchus which lays its eggs in the empty
shells of pelecypods such as Chione antigua (Cancelles 1944).
The commonest cormorant in the gulf is the King Cormorant P. albiventer^
60
which occurs nearly everywhere. Roosts of up to several hundred birds are
scattered along the coast, two of the largest being near Bahia San Roman.
The Blue-eyed Cormorant P. atriceps associates with King Cormorants,
comprising from 5 to 10 per cent of mixed flocks. These two species feed
together mainly in deeper waters. Blue-eyed Cormorants seem to be com-
moner near the mouth of the gulf than farther inshore.
One of the most striking aspects of cormorant biology was the rarity of
immatures among the non-resident species. Only 4 of 84 Rock Shags were
immature; immatures comprised no more than 10 per cent of the King
Cormorant flocks, and only one immature Blue-eyed Cormorant was identi-
fied, despite careful searching. Many young Olivaceous Cormorants were
present, but we made no estimates of their relative abundance.
Flamingos, Herons: Several hundred Chilean Flamingos Vhoenicopterus
chilensis, including one white-plumaged juvenile that appeared to be no more
than two-thirds grown, wintered along the southwestern corner of the gulf
in 1 97 1. Fewer were present in the same area in 1972. Whether the apparent
decrease reflects the later time of observation or local differences in the
ecology of nearby inland lakes where flamingos also feed, is unknown.
Human disturbance may also be important, for even in this remote area we
found several birds that appeared to have been shot.
In both years Common Egrets Casmerodius albus were rare, and were found
only on the south side of the gulf. Black-crowned Night Herons Nycticorax
nycticorax were commoner and are presumably resident. Although these
birds could be found on rocky outcrops along the entire coast, the majority
of the population was centered at a roost on Isla de los Pajaros. The extremely
dark coloration of these herons was striking. In 1971 Jehl saw two melanistic
birds there and several that approached this condition; in 1972 we also saw
several extremely dark individuals, but none that was truly melanistic.
Waterfowl: The main habitat for waterfowl is along the south and west
coasts where extensive clay terraces and mudflats are exposed at low tide.
The Crested Duck Lophonetta specularoides is by far the most abundant species,
occurring along all coasts. Isolated pairs are the rule, but flocks of up to ten
were seen. This species breeds locally, but migratory birds from farther
south probably augment the winter population.
Flying Steamer Ducks Tachyeres patacbonicus are probably winter visitors to
the area. In the gulf they are found almost exclusively along the west and
southwest shores. Small numbers of Brown Pintails Anas georgica and Yellow-
billed Teal Anas flavirostris were present, but these species prefer ponds
inland on the peninsula.
Sheld Geese Chloephaga spp. are seldom seen along the coast, but Golfo
San Jose seems to be on the migration path. On 12, 13, 14 and 19 August, 17
flocks of up to 200 geese, totalling 1,100 individuals, were observed heading
southwest over the gulf. Nearly all were Upland Geese C. picta, but two
Ruddy-headed Sheld Geese C. rubidiceps were identified among them. (These
migrants are not included in Table 1.). The increased numbers of Upland
Geese utilizing the shore of the gulf in 1972 as compared to 1971 is doubtless
due to the arrival of migrating individuals.
Hawks and Falcons: Except for a single Aplomado Falcon Y alco femoralis in
1 971 and an immature Peregrine Falcon F. peregrinus in 1972, we saw no
evidence of predatory birds along the shore of the gulf. The Aplomado
Falcon was hunting shorebirds at a mudflat; the Peregrine made a playful
61
pass at a small group of night herons. In 1972 we found a South American
Tern Sterna hirundinacea that had been eaten by a falcon.
Several large nests on steep cliffs along the south shore were presumably
made by Black-chested Buzzard-Eagles Geranoaetus melanoleucus, but we did
not observe this species in the immediate area.
Shorebirds: American Oystercatchers Haematopm palliatus are abundant
residents of Golfo San Jose. Along the sandy beaches of the south and east
coasts the population averages eight to ten pairs per mile. Cliffs along the
north shore limit oystercatcher habitat there, but even so the birds remain
extremely common. We estimated the population at 600 individuals. The
species seems to maintain territories all the year, so that in most areas pairs
are spaced out regularly along the beach. However, in some areas, such as
Isla de los Pajaros, flocks of 20 or more occurred and fed together without
interaction on the exposed mudflats.
Figure 3. A pair of American Oystercatchers Haematopus palliatus left, and a hybrid with
a Blackish Oystercatcher H. ater, Golfo San Jose.
In several locations (e.g., Punta Conos, Isla de los Pajaros) dark mudstones
outcrop along the bay, and many are covered with extensive mussel beds.
These areas provide habitats for the small resident population of Blackish
Oystercatchers H. ater. This species occasionally hybridizes with H. palliatus
in this area (Fig. 3, and Jehl in prep.).
Wintering Magellanic Oystercatchers H. leucopodus occupied the mudflats
of the southwestern corner in each year. These birds tended to remain
together in large flocks and did not usually associate with or interact with
the American Oystercatchers that utilized the same feeding areas.
Small flocks of Falkland Plovers Charadrius falklandicus occurred on almost
every mudflat. Most flocks consisted of 10 to 15 birds, though some were as
large as 200. The huge increase in 1972 is almost certainly a result of improved
censusing.
62
An unexpected find was that small numbers of Magellanic Plovers
Pluvianellus soda/is wintered in the area. This is the northernmost report for
the species. Details of the winter ecology and behaviour of this rare shore-
bird will be published elsewhere (Jehl in prep.).
The strongly compacted clays which form the beach over much of Golfo
San Jose are almost impenetrable. Consequently, feeding habitats for probing
birds are virtually lacking. Small numbers of ''summering" northern
hemisphere sandpipers Calidris canutus, C. bairdii, C. fuscicollis, C. alba were
found in each year. These birds tended to feed in intertidal regions in associa-
tion with Falkland Plovers.
Least Seedsnipe Thinocorus rumicivorus were seen sparingly in 1972 only.
Scattered individuals fed on exposed tideflats along the southwestern shore,
and small flocks were occasionally flushed from sand dune areas along the
eastern shore.
Snowy Sheathbills Chionis alba were found on clayflats and mussel beds
along the entire coast but principally on the south shore.
Skuas, Gulls, Terns: Approximately six Great Skuas Catharacta skua
wintered in the gulf in 1972 and were usually found in association with the
flock of Sooty Shearwaters. Most of the birds seen well, as well as two that
were collected, had reddish underparts and were referable to C. s. chilensis.
However, at least one (two?) bird in the gulf, seen on 9 and 16 August, was
extremely dark and certainly was not of that race. Two of the three Great
Skuas seen well in 1971 were also referable to chilensis. We watched two skuas
making alternate swoops on a swimming and apparently healthy Great
Grebe, but without success.
A population of 4,300 Kelp Gulls Icarus dominicanus was estimated in 1972.
As in 1 97 1, well over 95 per cent of the birds were adult. A small colony
breeds at Punta Conos, where 30-50 deserted nests were found, and a large
colony nests at Isla de los Pajaros (Ragonese & Piccinini 1972). Brown-
hooded Gulls L. maculipennis were far commoner in 1972 than in 1971,
perhaps because of an influx of migrants. The main concentrations occurred
along the south and east shores of the gulf. By mid- August many of the birds
were vocal and were engaging in apparent aerial territorial displays. Dolphin
Gulls Leucophaeus scoresbii were rare and we encountered them only in the
vicinity of cormorant or sea lion roosts, where they fed on scraps or on
faecal material.
South American Terns Sterna hirundinacea were uncommon in the gulf,
although flocks of up to 20 might be encountered. Royal Terns Thalasseus
maximus breed on the Valdes Peninsula (including Isla de los Pajaros;
Ragonese & Piccinini 1972; Figs. 18, 24), but virtually all depart the area in
winter; we saw only one bird in 1971 and four in 1972. A few Trudeau's
Terns Sterna trudeaui were found along the south shore of the gulf in 1972;
none were seen in 1971.
References :
Cancelles, A. 1944. Catalogo de los moluscos marinos de Puerto Quequen (Republica
Argentina). Rev. Mus. de la Plata {Nov. Ser.), Sec. Zoo/. 3 : 233-309.
Ragonese, A. E. & B. G. Piccinini. 1972. La vegetacion de la "Isla de los Pajaros" (Pro-
vincia de Chubut). Idia 291: 31-50. (Inst. Nacional de Tecnologia Agropecuaria,
Argentina).
Boswall, J. & R. J. Prytherch. 1972. Some notes on the birds of Punta Tombo, Argentina.
Bull. Brit. Orn. CI. 92: 1 18-129.
Zapata, A. 1967. Observaciones sobre aves de Puerto Deseado, Provincia de Santa Cruz.
Hornero 10: 351-378.
63
The birds of the Shira Plateau and west slope
of Kibo, Kilimanjaro
by David G. King
Received ioth January, 197)
INTRODUCTION
This paper is devoted to the birds of the Shira Plateau and the west slope of
the Kibo summit of Mount Kilimanjaro, Tanzania. An attempt is made to
list all species that have been observed in the area, and to add new information
on their distribution, status and habits.
The avifauna of the high altitudes of the south slope of Kilimanjaro has
been well studied and documented by a number of observers (Johnston 1886;
Oberholser 1905, 1906; Kittenberger 1958, 1959; Moreau 1935, 1944;
Moreau & Moreau 1939; Guest & Leedal 1954; Lamprey 1965). However,
the high altitudes of west Kilimanjaro have been largely neglected due to
their inaccessibility. The only workers to make observations in this area are
Swynnerton (1949) and Salt (1954, 1955) from a visit they made in November
1948, and Guest & Leedal (1954) from a visit in August 1953. Neither party,
spent much time in the area, so their observations were incomplete. In
October 1968 the problem of access was alleviated with the completion of a
track by the Tanzania Forest Service. This led to extensive exploration and
studies of the vegetation and fauna by the staff of the College of African
Wildlife Management at Mweka, near Moshi, and by others. This report
results from these expeditions.
The Shira Plateau is defined as the relatively level plain on the west
shoulder of Kilimanjaro between 11,000 and 12,500 feet contours, an area
of about 21 square miles. This is in contrast to the limits of 1 1,500 and 13,500
feet used by Salt (1955) which are not realistic nor in agreement with modern
topographical maps. All of the slope above 12,500 feet and between the
Machame Escarpment on the south and the Lent Group on the north is
considered the west slope of Kibo.
The Shira Plateau is surrounded on the west and south sides by the Shira
Ridge that rises to 13,000 feet. All streams that dissect the area merge and
flow away to the north-west. Eccentrically placed in the southern half of the
plateau is a large conspicuous cone shaped hill, Platz Kegel.
The dominating vegetation over most of the area from 11,000 to 13,500
feet is heath Erica, Philippia and Artemisia spp., interspersed with grasses
Deschampsia, Festuca and Pentaschistis spp., and everlastings Helichrysum spp.
However, the drainage lines and about 2 «5 square miles around the base of
Platz Kegel are open grasslands or bogs dominated by sedges, grasses and
herbs Ranunculus, Alchemilla and Lobelia spp. Above 13,500 feet West Kibo is
largely an alpine desert with few plants except in wet drainages; and above
1 5,500 feet only barren rock, scree, snow and ice.
This paper results from seven trips to the Shira Plateau and West Kibo
between October 1968 and September 1969. All parts of the study area were
explored and many parts were visited several times. It was only during trips
in July, August and September 1969 that bird specimens were collected.
These are now in the Mweka Wildlife College Museum except for six (two
Cisticola hunteri, one C. brunnescens, one Serinus striolatus, two Cercomela
sordida) donated to the British Museum (Natural History). Species of birds
that have been recorded in the alpine zone below 11,000 feet are mentioned
in a section 'Other species' towards the end of the paper.
64
I divided the birds into those that were clearly resident in the Shira area
and into those that were occasional or rare visitors. Three species, namely
Mountain Buzzard Buteo oreophilus, Stonechat Saxicola torquata and Lammer-
geyer Gypaetus barbatus, listed as residents of high elevations by Lamprey
(1965) and Moreau (1944, 1966), are included here only as visitors. My list
includes two species, Apus aequatorialis and Cisticola brunnescens, not previously
recorded from upper Kilimanjaro.
All specimens collected were compared with those from other parts of
Kilimanjaro that are either in the Mweka Wildlife College Museum, the
National Museum of Kenya or the British Museum (Natural History). I have
followed the systematics used by White (1960-62, 1961-63, 1965), although
among others, Mackworth-Praed & Grant (i960), Blake & Vaurie (1962),
Ripley (1964) and Howell et al. (1968) were consulted.
RESIDENT SPECIES
Augur Buzzard, Buteo rufofuscus augur Rupp. : Two individuals were seen
on almost every trip to the Shira Plateau. This species had also been noted by
both Swynnerton (1949) and Guest & Leedal (1954). All sightings were
likely of the same two birds as one was of the typical white phase and the
other of the melanistic phase. Moreover, they were frequently together. They
roam over the entire plateau and ascend West Kibo to at least 14,000 feet.
On one occasion I observed the white individual with a rat or mouse. Rodent
probably form the bulk of the augur's diet as they are abundant and no
resident game birds are known above 10,000 feet.
Swifts, Apus melba africanus Temminck and A. ae. aequatorialis (von
Muller): Large swifts are seen regularly on the south slope of Kilimanjaro
to 17,000 feet. Lack (in Moreau 1936) identified swifts he saw at Horombo
Hut as Apus melba but Kittenberger (1959) those he saw on Kibo as the
similar species, A. aequatorialis. However, there is no record of a specimen
of either species having been collected from high elevations. Sjostedt (in
Friedmann 1930) collected A. melba in 1905 above Kibongoto (4,400 feet) on
south Kilimanjaro, which is the only specimen of the large swifts from the
mountain. On Mount Meru, 40 miles to the west, I collected an individual of
A. aequatorialis at 8,300 feet. I feel certain that swifts I saw swooping and
diving over the Shira Plateau on 7th September were A, aequatorialis ', as their
mottled bellies were clearly visible. In summary, it is probable that both
species are resident, although confirmation from specimens is needed. Swifts
do not appear to be as numerous in the Shira area as elsewhere on Kiliman-
jaro, as I saw them only two or three times. Swynnerton (1949) recorded
them only once.
Hill Chat, Cercomela sordida hypospodia (Shell.): This chat is the most
common and widespread bird found on upper Kilimanjaro. It prefers the
areas of broken heath and of grassland interspersed with everlastings and low
heath which are the common habitats of the Shira Plateau. It is less common
in the denser bush below 1 1,000 feet, that is, the edge of the plateau; and in
areas of dense heath on the plateau. It also becomes scarce above 13,500 feet
although occurring to the highest vegetation.
I collected four specimens on 3rd-5th July, four on 10th August and two
on 6th September. The July birds were not sexed. The August collection
was of three males and a female, and the September group of two males.
None of the August birds were in breeding condition but the testes of both
65
September males were greatly enlarged. Moreover, in September the birds
were dispersed rather than in the flocks of five to eight that are seen through
most of the year.
On 7th September 1969, my companions V. C. Gilbert and M. Bigger
discovered a nest at 13,000 feet on a south facing slope of a stream. The nest
was a grass cup at the end of a 1 2-inch deep cleft between two rocks and
under a third. It contained two eggs and a newly hatched chick and shell.
The eggs were thrush blue (turquoise blue) with small blotches the colour
of dried blood on the large end. They were approximately 23 by 17 mm. The
only other breeding record I could find for C. s. hypospodia is that given by
Mackworth-Praed & Grant (i960). They record breeding in June (source
not given) and describe the eggs as "white with faint tawny spots and
streaks". The eggs found by Gilbert and Bigger are more similar to those
of C. s. ernesti (Sharpe) of Mount Kenya (Mackworth-Praed & Grant 1 960)
although it is well known that variation does occur in the eggs of chats.
Hunter's Cisticola, Cisticola hunteri hunteri Shell. : This warbler occurs in
the heath of Kilimanjaro to about 13,500 feet and is the second most com-
mon species on the Shira Plateau. Lynes (1930) and Mackworth-Praed &
Grant (i960) say it is normally associated with dense shrubby undergrowth,
but on the plateau small flocks of six to ten birds were frequently seen in
moorlands with only the sparest of shrubby cover. They associate with the
Hill Chat in flocks, and the two species appear to be competing directly for
food, that is, insects. At elevations below 11,000 feet, however, the warbler
is the dominant species.
I collected 12 specimens at approximately 12,000 feet: three unsexed
specimens on 5 th July, three males and two females on 10th August, and
three males and a female on 6th September. None were in breeding condition.
All were similar in size and colour to those in the museums collected above
treeline since 1889 (when Hunter made the first collection).
Several authors including Oberholser (1905), Lynes (1930), Moreau &
Moreau (1939) and White (1962) noted that specimens of Hunter's Cisticola
are more sombre at high elevations than at lower elevations on Kilimanjaro.
Specimens from under 9, 5 00 feet have more russet on the head and thighs,
and their general coloration is more brown than grey. Moreover, the chin
is whitish rather than grey. Specimens I examined from all elevations of
Kilimanjaro were darker than specimens from all elevations on Mount Meru,
Mount Kenya, the Aberdares and all other localities, although included in the
same subspecies, C. h. hunteri^ by White (1962). Prior to White (1962)
specimens from all these localities except Kilimanjaro were called C. h.
prinioides Neumann (Mackworth-Praed & Grant i960) because of the colour
difference. I also noted that some rectrices of most specimens from upper
Kilimanjaro had many conspicuous "feather checks" but were uncommon
on specimens from other localities.
White-naped Raven, Corvus albicollis Lath.: On all trips to the Shira
Plateau two individuals, presumably a pair, were observed. They occasionally
ascend the west slope of Kibo to at least 17,000 feet where I saw them flying
high over the glaciers. The raven is relatively common over the entire
mountain and has been observed as high as the summit of Kibo (19,340 feet)
many times.
Scarlet-tufted Malachite Sunbird, Nectarinia johnstoni johnstoni Shell. : This
sunbird is numerous at high elevations wherever lobelias Lobelia deckenii are
66
in flower. They appear to rely largely on the nectar of lobelia for food —
although I have seen individuals pursuing insects — and seem to move about
the mountain seeking them. In April 1969, these sunbirds were common in
the valleys of West Kibo between 12,500 and 14,000 feet where the lobelias
were in flower. In early July lobelias and sunbirds were very common near
Platz Kegel but on 10th August only a single individual was seen in the same
area. The lobelias had finished flowering and the birds had gone elsewhere in
search of food. This is probably why Williams, in July 1949, encountered few
of this species on south-east Kilimanjaro (Williams 195 1).
One male was collected on 5 th July, but the size of the testes was not noted.
Its measurements were within the limits given by Williams (195 1) for N.j.
johnstoni.
Yellow-crowned Canary, Serinus canicollis flavivertex (Blanf.) : This small
canary is occasionally seen on the Shira Plateau and the west slope of Kibo
to 13,000 feet. It usually occurs in small flocks in the more open heath and
grasslands, but as it is a wary bird, it is seldom seen close at hand. On 4th July
I saw a male courting a female near giant groundsels Senecio cottonii in a valley
west of Platz Kegel. As the female moved slowly from limb to limb the male
kept approaching with much fluttering of the wings and excited chirping.
The female eventually appeared to tire of his pursuit and flew farther away.
The male burst into song before I collected him. His testes were greatly
enlarged. This single specimen was indistinguishable from a male collected
on 10th August at 6,000 feet on west Kilimanjaro. This latter bird was also
in breeding condition.
Streaky Seed-eater, Serinus striolatus subsp. : Lamprey (1965) says that the
Streaky Seed-eater is "A common species which occurs in flocks on the
moorlands up to 14,000 feet". I have seen few of this finch on the Shira
Plateau or elsewhere on Kilimanjaro, nor is it recorded as being "common"
by any other observer. A single individual seen sitting on a giant groundsel
5 th July was collected. One of two seen with a flock of Hunter's Cisticolas
near Platz Kegel 10th August was also collected. The first specimen was not
sexed but the second was a male in non-breeding condition. I cannot find
any previous record of this species having been collected above the forest of
Kilimanjaro.
The Shira specimens were compared with those in the National Museum
of Kenya, British Museum (Natural History) (including eight from lower
Kilimanjaro), and Mweka Wildlife College Museum, and were found to
differ from all of them. The distinguishing feature of the Shira specimens is a
very conspicuous lemon-yellow wash throughout the plumage. This in-
cludes the superciliary stripe and the light markings on the sides of the head
and underparts from the chin to the vent. The yellow was very noticeable
even in the field. Of the large collection of Streaky Seed-eaters in the National
Museum of Kenya only three females from Kinankop, Aberdares, have a
definite yellowish wash, and this is very light and confined to the underparts.
Specimens in the British Museum from the moorlands of Mount Kenya and
elevations below 7,000 feet on Kilimanjaro have a slight wash on the head
and throat, but none approach the specimens from upper Kilimanjaro for
brilliance of colour. The difference may be sufficient for naming of a new
subspecies; however, additional specimens are needed. The form that lives
on lower Kilimanjaro is somewhat in doubt as White (1963), evidently
following Sclater (in Moreau 1935), calls it S. s. striolatus ', but Howell et al.
67
(1968) use S. s. affinis, a name first proposed by Richmond (Oberholser
1905).
With further reference to the Shira specimens, there are two in the British
Museum that should be noted. They are a bird collected by Moreau in 1934
in the Mbulu District of northern Tanzania and another from Ruwenzori,
both strongly coloured but with orange-buff, not lemon-yellow. C. H. B.
Grant marked the label of the Mbulu specimen as juvenile, and the other
almost certainly is as well. The Shira specimens may also be juveniles.
OCCASIONAL AND RARE VISITORS
Lammergeyer, Gypaetus barbatus meridionalis Keys. & Bias.: This very
large bird of prey is rarely seen on Kilimanjaro and its status as a resident is
in question (Guest & Leedal 1954). I spent a total of nearly 60 days above the
forest between October 1968 and September 1969, and only saw a Lammer-
geyer once. On 6th November 1968, G. S. Child and I saw an individual
circling over the Shira Plateau. I am unaware of any other recent sightings of
the species from Kilimanjaro, so it may be worthwhile to note an individual
that I saw at 13,000 feet on Mount Meru on 29th September 1968.
Mountain Buzzard, Buteo oreophilus oreophilus Hart. & Neum. : Although
recorded in large numbers from the upper slopes of Kilimanjaro by Moreau
(1935), I have only seen this buzzard on the Shira once. This was on 7th
September 1969, when a lone individual flew by in a southerly direction. I
have not seen this species above the forest elsewhere on Kilimanjaro, nor
does it seem as numerous in the forest as in the same zone on Mount Kenya
(pers. obs.).
Quail, Coturnix coturnix erlangeri Zedl. : This species was first recorded
above the forest of Kilimanjaro by Lamprey (1965). Child saw a single
individual near our camp on the Shira Plateau on 3rd July which he duly
collected. This bird, a female, was on a very open ridge sparsely covered
with grasses and low heath. The ovary was regressed and the crop was filled
with unidentified seeds. This bird was likely a stray from large numbers that
were seen during the same trip in Londrossi Glade, west Kilimanjaro, at
7,000 feet. On 7th September 1969, I saw two more quails on grassy moor-
lands near Platz Kegel. It is doubtful that this species is resident and I believe
all sightings to be of birds on migration.
The subspecific name erlangeri is used, following Benson & Irwin (1966),
who applied this name to the populations of eastern Africa, and restricted
C. c. ajricana Temminck & Schlegel to those breeding in southern Africa
south of the Limpopo.
Plover sp. : Swynnerton (1949) reported sighting a plover in the boggy
valley west of Platz Kegel. He believed it was a species of Stephanibyx. On
7th September 1969, 1 saw a large wader in a stream which I was not able to
identify positively. However, it definitely was not a Stephanibyx sp. as it
lacked a black tail band. The bird was Stephanibyx in size with white under-
pays and brownish uppers. Possibly it was a Greenshank Tringa nebularia on
migration.
African Snipe, Gallinago nigripennis nigripennis Bonap. : This wading bird
was recorded in November 1948 by Swynnerton (1949) from the boggy area
near Platz Kegel. I searched the same area in November 1968 and in July,
68
August ami September 1969, but failed to find more of this species. M.
Bigger (pers. comm.) saw an individual in the bog on 17th July 1965, and in
January 1970 V. C. Gilbert (pers. comm.) saw two. These sightings suggest
snipe are regular visitors to the area.
Owl sp. : An owl or eagle-owl was recorded by Guest & Leedal (1954)
near the Lent Group on West Kibo and at the base of Mawenzi Peak.
I have not observed or heard any species of owl or eagle-owl in the Shira
area and neither did Swynnerton (1949). However, the presence of a species
could be expected as there is a very high rodent population in the area and
an eagle-owl Bubo capensis is known to reside at high elevations on Mount
Kenya. C. W. Benson (pers. comm., Jan. 1970) suggests the bird recorded by
Guest & Leedal may have been this species; also, see Benson & Irwin (1967).
Stonechat, Saxicola torquata axillaris (Shell.) : This chat is common in the
heath just above the forest. On 6th September 1969, I observed a lone indi-
vidual near the centre of the Shira Plateau at 11,700 feet. I have several
records and one specimen from the heath at 11,000 feet at the edge of the
plateau. However, I believe it is only an occasional visitor out onto the
plateau proper, and therefore have not included it in the list of residents
above.
Pectoral-patch Cisticola, Cisticola brunnescens subsp. : On 7th September
1969, 1 saw four or five small cisticolas at 1 1,800 feet on an open moor north
of Platz Kegel. I collected two, both females, one of which was taken to the
British Museum. There it was tentatively identified by Mrs. B. P. Hall as
Cisticola brunnescens. Mrs. Hall said (pers. comm., Jan. 1970) the specimen
appeared to belong to a population intermediate between C. b. hindii Sharpe
of Kenya and C. b. cinnamomea Reichw. of the south. In the British Museum
there is a female from Engare Nairobi (5,400 feet) on the west side of Kili-
manjaro, and another female from 6,500 feet on Mount Hanang, having the
same characteristics. Mrs. Hall said that three more similar specimens from
the Masai Steppe are in Berlin. Lynes (1930), however, included them in C. b.
hindii without comment. All in all, very little is known about the distribution
or habits of these birds and they could represent an as yet undescribed sub-
species of C. brunnescens. It is also possible that the species is resident on the
Shira Plateau grasslands even though not previously recorded. An interesting
feature of both specimens was the presence of several toes lacking claws and
being swollen at the tips.
Swallow sp. : Two small dark swallows were seen on 5 th July flying over
the south rim of the plateau. They may have been Psalidoprocne bolomelaena y
which is usually found in the forest belt below 9,000 feet. I have seen swallows
above the forest of Kilimanjaro on other occasions and as high as 14,000
feet on Mount Kenya.
Dioch spp. : The remains of two different Que lea spp. were recorded by
Guest & Leedal (1954) high on Kilimanjaro. One of these was on West Kibo
but it is thought that both were accidental casualties during migration. There
are no recent records.
OTHER SPECIES
There are a number of species which have not been recorded at high
elevations on Mount Kilimanjaro but should be mentioned. These include
69
species that have been recorded between treeline and 1 1 ,000 feet or at high
elevations elsewhere in East Africa, particularly Mount Kenya, On 5th-7th
July 1969, a Fiscal Shrike Lanius collaris was seen several times at 10,500 feet
on west Kilimanjaro. At the same locality on 10th August, I collected a
Pangani Longclaw Macronyx aurantiigula. On Mount Kenya the African
Black Duck Anas sparsa breeds in the moorlands but has not yet been ob-
served above 8,000 feet on Kilimanjaro. The Mountain Francolin Franco linus
psilolaemus, which breeds in the moorlands of Mount Kenya, does not occur
on Kilimanjaro, but the closely related Shelley's Francolin F. shelleyi has
been seen up to 10,000 feet (Lamprey 1965). It may go higher. Two residents
of the alpine zone of Mount Kenya, the Slender-billed Starling Onychognathus
tenuirostris and the rarely seen Scarce Swift Apus myoptilus, are unknown on
Kilimanjaro, possibly because their ranges do not include this mountain. It
is perhaps surprising that more birds of prey have not been recorded. At least
three harriers Circus spp., the Black Kite Milvus migrans^ and Verreaux's
Eagle Aquila verreauxii have been observed over the moorlands of the Aber-
dares or Mount Kenya.
Undoubtedly over the next few years more species will be added to those
already recorded in the higher altitudes of Kilimanjaro. Access is steadily
improving and now that the mountain has been made into a national park
even greater numbers of visitors can be expected. It has been the objective
of this paper to bring together all that is known of the bird fauna of upper
Kilimanjaro, particularly the less well known western flank, and provide a
base for future observers to build on.
AC KNOWLEDGEMENTS
I am particularly grateful to the Mweka College of African Wildlife
Management which provided the opportunity and means to explore the
Shira Plateau and west slope of Kibo; and which provided laboratory space
and technical assistants. I also wish to extend warmest thanks to the many
individuals who assisted in the field work, particularly Gil Child and V. C.
"Tom" Gilbert. In addition, without the co-operation of the National
Museum of Kenya and of the British Museum (Natural History), including in
the latter Dr. D. W. Snow, D. Goodwin and Mrs. B. P. Hall, this paper could
not have been written.
SUMMARY
An account is given of the birds of the Shira Plateau and the west slope of
the Kibo summit, upper Kilimanjaro. Data include a breeding record for a
chat Cercomela sordida and the dependence of a sunbird Nectarinia johnstoni
on flowering Lobelia deckenii. The populations of a seed-eater Serinus striolatus
and a cisticola Cisticola hunteri appear to differ in colour of plumage from those
at lower altitudes on the mountain. Apus aequatorialis and Cisticola brunnescens
are recorded from upper Kilimanjaro for the first time, and first specimens of
the latter and Serinus striolatus were collected.
POSTSCRIPT
C. W. Benson (pers. comm.) suggests that the Chestnut-tailed Pygmy
Crake Sarothrura affinis could occur on upper Kilimanjaro. It has been col-
lected in the moorland zone of Mount Kenya and the Aberdares between
11,000 and 12,000 feet, and its occurrence to the southward of Kilimanjaro
is well established (Keith et al., 1970, Bull. Amer. Mus. Nat. Hist. 143(1):
5 7-59)-
70
The author made another visit to upper Kilimanjaro from ioth to 14th
February 1973, in the hope of obtaining more data on the Streaky Seed-eater
and Pectoral-patch Cisticola. This was prevented by poor weather; however,
good sight records were had of the Alpine Swift Apus melba at 1 6,000 feet
and the Abyssinian Nightjar Caprimulgus poliocephalus at 10,500 feet. The
latter species had not previously been recorded from Kilimanjaro, although
known from Mount Kenya and the Aberdares. It was giving a very loud
call of pure flute-like notes that sounded like "peeyou" followed by "pee-
whuwhuwhu", this latter phrase descending. This call had been heard in
1968-69, but the bird could not be identified, as calling is only heard in the
late evening and after dark, and only from the ground or low plants such as
Lobelia.
References :
Benson, C. W. & Irwin, M. P. Stuart. 1966. The Common Quail Coturnix coturnix in the
Ethiopian and Malagasy Regions. Arnoldia (Rhodesia) 2 (13): 14pp.
— 1967. The distribution and systematics of Bubo capensis Smith (Aves). Arnoldia (Rhodesia)
3(19): 19pp.
Blake, E. R. & Vaurie, C. 1962. Family Corvidae, Crows and Jays. In Check-list of birds of
the world. 15: 204-282. Cambridge, Mass.: Mus. Comp. Zool.
Friedmann, H. 1930. Birds collected by the Childs Frick Expedition to Ethiopia and Kenya
Colony. 1. Non-passeres. U.S. Nat. Mus. Bull. 153.
Guest, N. J. & Leedal, G. P. 1954. Notes on the fauna of Kilimanjaro. Tang. Notes & Recs.
36: 43-49-
Howell, T. R., Paynter, R. A. & Rand, A. L. 1968. Family Fringillidae : Subfamily Car-
duelinae, Serins . . . and allies. In Check-list of birds of the world. 14: 207-306. Cambridge,
Mass.: Mus. Comp. Zool.
Johnston, H. H. 1886. The Kilimanjaro Expedition. London: Kegan Paul, Trench & Co.
Kittenberger, K. 1958. My ornithological collecting expeditions in East Africa. Part 1.
Aquila 65 : 13-37.
— 1959. My ornithological collecting expeditions in East Africa. Part 2. Aquila 66: 53-87.
Lamprey, H. F. 1965. Birds of the forest and alpine zones of Kilimanjaro. Tang. Notes &
Recs. 64: 69-76.
Lynes, H. 1930. Review of the genus Cisticola. Ibis (12)6, suppl.
Mackworth-Praed, C. W. & Grant, C. H. B. i960. African handbook of birds. Series I, Vols.
1 & 2. 2nd. ed. London: Longmans.
Moreau, R. E. 1935. A contribution of the ornithology of Kilimanjaro and Mount Meru.
Proc. Zool. Soc. Lond.: 843-891.
— 1944. Kilimanjaro and Mount Kenya: some comparisons, with special reference to the
mammals and birds; and with a note on Mount Meru. Tang. Notes & Recs. 18: 28-59.
— 1966. The bird faunas of Africa and its islands. London: Academic Press.
Moreau, R. E. & Moreau, W. M. 1939. A supplementary contribution to the ornithology
of Kilimanjaro. Rev. Zool. Bot. Afr. 33: 1-15.
Oberholser, H. C. 1905. Birds collected by Dr. W. L. Abbott in the Kilimanjaro Region,
East Africa. Proc. U.S. Nat. Mus. 28: 823-936.
— 1906. Notes on birds from German and British East Africa. Proc. U.S. Nat. Mus. 30:
801-811.
Ripley, S. D. 1964. Family Muscicapidae : Subfamily Turdinae, Thrushes. In Check-list of
birds of the world. 10: 13-272. Cambridge, Mass.: Mus. Comp. Zool.
Salt, G. 1954. A contribution to the ecology of upper Kilimajaro. /. Ecol. 42: 375-423.
— 1955. The Shira Plateau of Kilimanjaro. Geogr. Journ. 117: 150-164.
Swynnerton, G. H. 1949. Report on a safari to explore the Shira Plateau, west Kilimanjaro,
Ann. Rep. Game Dept. Tang., 1948: 9-1 1.
White, C. M. N. 1960-62. A check list of the Ethiopian Muscicapidae (Sylviinae). Occ. Pap.
Nat. Mus. S. Rhod. 24B: 399-430; 26B: 653-738.
— 1961-63. Revised check lists of African passerine birds. Lusaka: Government Printer.
— 1965. A revised check list of African non-passerine birds. Lusaka: Government Printer.
Williams, J. G. 195 1. Nectarinia johnstoni: a revision of the species, together with data on
plumages, moults and habits. Ibis 93: 579-595.
7i
Remarks on the status of Campethera "scriptoricauda"
and related species
by Lester L. Short
Received 26th February, 197)
In reviewing the genus Campethera as part of my preparation of a monograph
of the Picidae, I examined specimens of the superspecies C. nubica, including
C. punctuligera, C. nubica, C. bennettii, and C. "scriptoricauda". I was able to
examine only eleven specimens of the last-mentioned form, but some addi-
tional features were found to shed light on the relationships of scriptoricauda.
Various authors have recognized the very close relationship of the putative
species comprising this complex (see, e.g., Chapin 1939:365; Clancey 1964:
162; Goodwin 1968:21). The allospecies (Amadon 1966) nubica, bennettii and
punctuligera are closely similar and replace each other geographically without
overlapping, as far as is known; it is entirely possible that they are conspecific.
Compared with these three forms, scriptoricauda, allopatric with, and occur-
ring between the ranges of bennettii and nubica, is less distinctive, and in fact
this taxon presents taxonomic difficulty because it combines features of
nubica and bennettii, while lacking characteristics that are not shared by those
two forms.
Comparing scriptoricauda with nubica, similarity is shown in: spotting of the
throat (found in scriptoricauda and in a few specimens of nubica, although 10 of
16 birds from Dire Daoua, Boule Boule, Lake Zwai and Maraco, Ethiopia,
have the throat and chin partly, or in 3 cases, fully spotted); streaked ear
coverts ; and the white background (both sexes) of the ear coverts, and the
throat and chin. One possibly distinctive feature of scriptoricauda is its yellow-
based black bill, but dried specimens of bennettii resemble scriptoricauda in bill
colour (some specimens of bennettii bear labels having bill with a "pale base"
as do several specimens of scriptoricauda at my disposal). Until we know more
precisely the variation in bill colour of bennettii this trait of scriptoricauda
cannot be appraised. In wing length and in tail length scriptoricauda falls
within the upper range of variation of nubica, and in the low range (wing to
122 mm, Belcher 1928) of bennettii. Resemblances between scriptoricauda and
bennettii are: back coloured yellow-brown (rather than green-brown of
nubica)\ back barred with white (rather than spot-barred as nubica); tail
barring obscure or vague (not crisply barred as in nubica); culmen of bill
curved (straighter in nubica); and, bill narrow across the nostrils. Regarding
this last feature, 11 specimens of scriptoricauda measure 2-6, 2-6, 2-7, 2-8,
2-8, 2-8, 2-9, 2-9, 2-9, 3-1 and 3-1 mm across the culmen between nostrils;
33 bennettii measure from 2-5 to 3-3 mm, and 5 5 nubica measure 3-2 to 4-2 mm.
(I note that related punctuligera is like bennettii and scriptoricauda, and not like
nubica in this measurement). The bill differences between bennettii and nubica
suggest that the latter uses the bill for tapping and excavating in the bark of
trees to a greater extent than does bennettii, and the resemblance of scriptori-
cauda to bennettii in bill structure suggests agreement with it rather than with
nubica in foraging habits. To these similarities must be added the vocal
resemblance of scriptoricauda to bennettii, as opposed to nubica (Benson 1948 : 61).
The range of scriptoricauda extends from central eastern Tanzania (70
miles north of Morogoro) to the northeastern corner of Mozambique, and
thence westward around the southern end of Lake Nyasa to southern
Malawi (White 1965; Clancey 1969b; specimen in Los Angeles County
Museum). C. b. bennettii occurs southwest of the range of scriptoricauda in
72
Mozambique, and northwest of it in northern Malawi and western Tanzania.
C. nubica "pallida" occurs in northeastern Tanzania, meeting scriptoricauda at
least sporadically (seasonal wanderers ?) at or near Morogoro. No interactions
with C. nubica have been reported ; hybrids of scriptoricauda and nubica would
be very difficult to detect. C. b. bennettii and scriptoricauda undoubtedly meet
in south-central Malawi. Benson (1952: 152) described intermediates of
bennettii and scriptoricauda from : 60 miles north of Tete (type of C. b. vincenti
Grant & Mackworth-Praed 1953), female, with unspotted, pinkish brown
throat and similarly coloured ear coverts ; female from Zobue with unspotted,
even paler (faintly buffy) throat and ear coverts; females from Kapeni-
Matindi, near Blantyre (bird with small throat spots on a beige-tinted back-
ground, noted as scriptoricauda approaching bennettii by Goodwin 1968:
20), and 30 miles south of Chikwawa, all with spots on chin and throat; and
males from Zomba, 60 miles north of Tete, and Kapiriuta, all with ear
coverts intermediate between the white of bennettii and the black-streaked
coverts of scriptoricauda, but without spots on the throat and chin. Goodwin
(1968) commented upon the lack of overlap between phenotypically pure
specimens of bennettii and scriptoricauda. The extent to which hybridization or
intergradation occurs is not known, but it is clear that C. bennettii morpho-
logically converges upon, rather than diverges from, scriptoricauda where
their ranges approach. I note that all but three of 1 1 specimens of scriptori-
cauda I have examined show red-brown colour, like that of bennettii in tone
(not dark brown as in nubica) , in a small area of the ear coverts behind and over
the eyes; one male has this brown colour suffused throughout the anterior
ear coverts.
A factor relating to an appraisal of scriptoricauda is that the variation found
in C. bennettii is extensive, as has been documented by Clancey (1964, 1967,
1968, 1969a) and by Irwin & Benson (1966). Particularly notable is the
variation in brown throat and ear covert colour among females, and its
presence among scattered males from throughout that species' range. Three
of nine immature males and five of 20 adult males representing C. b. bennettii
and C. b. capricorni have a light cinnamon brown throat, and generally
browner ear coverts. Further, although females show some clinal variation
from nearly black (capricorni) to pale brown (bennettii, "vincenti") ear coverts
and throat, some capricorni specimens are red-brown in those areas, and
bennettii specimens vary from cinnamon to nearly black. The possibility, even
the likelihood, of migration among some populations of C. bennettii (Clancey
1964: 164) suggests that there is extensive gene flow, perhaps accounting
for some of the observed variation. In any case, variation in colour of the ear
coverts and throat in bennettii, and of throat spotting in nubica indicates genetic
instability and plasticity relating to these features. It is unwise to assign
crucial importance to them in evaluating the status of scriptoricauda. I might
note that I fail to see, amid the variation in C. bennettii, a need for nomencla-
tural recognition of more than two subspecies, bennettii and capricorni, in
addition to scriptoricauda.
It seems most unlikely that scriptoricauda, which does not approach the
morphological distinctiveness of punctuligera , bennettii and nubica, will prove
reproductively isolated, and hence specifically distinct from bennettii. The
resemblance of scriptoricauda to bennettii in bill structure presumably corre-
lated with feeding habits, their vocal similarity, and their resemblance in
general features of plumage colour (back colour and markings, tail pattern)
as contrasted with more variable ear-covert and throat-chin patterns, incline
73
me to agree with Benson (1952: 153), White (1965) and others merging
scriptoricauda in bennettii. Obviously we need further data, especially from
areas where the various forms of this complex may meet. At this time,
however, although not discounting the possibility that the entire complex
may prove to represent a single biological species, the preponderance of
available evidence suggests that scriptoricauda is conspecific with bennettii. C.
punctuligera may prove conspecific with C. bennettii, which it resembles in bill
structure, and C. nubica, the most distinctive of these forms, is most apt to
prove specifically distinct from the others. For the time being I prefer to treat
punctuligera, bennettii and nubica as separate species, listed in that linear order.
As a final comment I suggest that C. abingoni (and related C. notata) may
have played a role in the evolution of the C. nubica complex. I believe
abingoni and notata are closely related to the nubica group. They are broader
billed, and generally perform more tapping than do species of the nubica
complex. The essential point is that broad-billed C. abingoni overlaps in range
with narrow-billed C. bennettii and punctuligera, but is essentially allopatric
with C. nubica, which has a relatively broad bill (morphologically overlapping
somewhat with abingoni). Thus competitive interaction with abingoni may be a
factor influencing the narrow bill and feeding habits (e.g., ground feeding
has been reported for bennettii and punctuligera, but not for nubica) of punctu-
ligera and bennettii ', whereas nubica ecologically may replace both the strong-
billed abingoni and its own allospecies punctuligera and bennettii.
Acknowledgements: I thank M. A. Traylor and K. Stager for the loan of
specimens vital to this study, respectively from the Field Museum of Natural
History, Chicago, and the Los Angeles County Museum. Other specimens
were available in the American Museum of Natural History.
Summary: Campethera "scriptoricauda" is found to agree with C. bennettii
and not nubica in bill shape and bill width, and in several colour characters,
as well as in voice and habits, as noted by other authors. It is treated herein
as a race of C. bennettii. The three woodpeckers C. bennettii, C. punctuligera
and C. nubica are very closely related, forming a superspecies. Competition
with another relative, C. abingoni, with which they are sympatric, may have
caused divergence of C. bennettii and punctuligera from C. nubica, which is
largely or completely allopatric with those three species.
References :
Amadon, D. 1966. The superspecies concept. Syst. Zool. 15 : 245-249.
Belcher, C. F. 1928. Notes on Nyasaland birds. 1. Additions to the list of Nyasaland birds
(Non-Passeres). Ibis (12) 4: 1-9.
Benson, C. W. 1948. Geographical voice variation in African birds. Ibis 90: 48-71.
— 1952. Notes from Nyasaland. Ostrich 23: 144-159.
Chapin, J. P. 1939. The birds of the Belgian Congo. Bull. Amer. Mus. Nat. Hist. 75 : 1-632.
Clancey, P. A. 1964. Miscellaneous taxonomic notes on African birds. XXII. Durban
Museum Novit 7 : 157-187.
— 1967. Critical comments on the subspecies of some birds from Zambia. Durban Mus.
Novit. 8: 77-107 .
— 1968. Subspeciation in some birds from Rhodesia. Durban Mus. Novit. 8: 11 5-1 5 2.
— 1969a. Miscellaneous notes on African birds. XXVII. Durban Mus. Novit. 8: 227-274.
— 1969b. A catalogue of birds of the South African sub-region. Suppl. no. 1. Durban
Mus. Novit. 8: 275-324.
Goodwin, D. 1968. Notes on woodpeckers (Picidae). Bull. Brit. Mus. (Nat. Hist.) 17: 1-44.
Grant, C. H. B. & Mackworth-Praed, C. W. 1953. A new race of woodpecker from
Portuguese East Africa. Bull. Brit. Orn. CI. 73 : 55-56.
Irwin, M. P. Stuart & Benson, C. W. 1966. Notes on the birds of Zambia: Part II. Arnoldia
(Rhodesia) 2: 1-2 1.
White, C. M. N. 1965. A revised check-list of African non-passerine birds. Lusaka: Government
Printer.
74
The Relationships of the Swallow-tanager Tersina viridis
by Charles G. Sibley
Received ntb April, 197)
INTRODUCTION
The Swallow-tanager occurs in tropical and lower subtropical areas from
Panama and Trinidad to Bolivia, southeastern Brazil, and northeastern
Argentina. It is a nine-primaried oscine bird with a relatively broad, flat bill
and long swallow-like wings. The adult male is mainly iridescent bluish or
bluish-green with the facial area, throat and flank bars black and the abdomen
white. The female is green with the face and throat greyish-brown and the
abdomen and flank bars pale yellow-green. Swallow-tanagers are territorial
when breeding but gregarious otherwise. They eat fruit and insects taken in
flight. The nest is built in a hole, sometimes excavated by the birds in a
vertical bank. Like many hole-nesting birds the eggs are pure white (Schaefer
1953)-
TAXONOMIC HISTORY AND CHARACTERS
The Swallow-tanager came to the attention of ornithologists early in the
last century and was at first assigned to the swallows because of the long
wings and broad bill. During the 19th century the Swallow-tanager was
placed in several different genera, most often in Procnias. This generic name
was later shown to belong to the bellbirds of the family Cotingidae (Ridgway
1907: 880) and since 1907 the Swallow-tanager has usually been placed in
Tersina of Vieillot (1 8 1 9).
It was recognized quite early in its taxonomic history that Tersina is
related to the tanagers. For example, Sclater (1862) placed it in its own
subfamily (Procniatinae) in the tanager family "Tanagridae" (=Thraupidae).
In 1895 Ridgway proposed the establishment of the family Procniatidae
based upon "the notable characters presented in the skull" which had been
called to his attention by F. A. Lucas. Lucas (1895a) published a brief note
announcing Ridgway's new family and (1895b) a description of the "charac-
ters of the Procniatidae."
Lucas (1895b) found that Tersina is like the tanagers in many characters but
that in its palatal skeleton it "departs so widely, not only from the tanagers,
but from the large majority of passerine birds, as to warrant the establishment
of a separate family for . . . the genus." The palatal differences involve "the
total absence of the transpalatine processes, the small size of the inter-
palatines, and the slenderness and outward curvature of the prepalatine
bars, which makes the interpalatine vacuity almost oval in shape." Lucas
noted that the absence of the transpalatine process was unique among the
passerines although found in Thinocorus and Turnix. Except for the palate, the
skull of Tersina was found by Lucas to be more like that of a tanager than
like that of a swallow. In its pterylosis Tersina was found to be either dis-
tinctive or similar to the tanagers. The shape of the dorsal tract in Tersina
resembles that of Thraupis palm arum although differing in size.
In a study of the hyoidean structures in passerine birds George (1962)
found that, with two exceptions, the New World nine-primaried oscines
have a laterally compressed basihyale. In Peucedramus and Tersina this bone is
cylindrical in shape. George therefore excluded the Tersinidae from the
nine-primaried assemblage although the Thraupidae were included.
Since 1895 Tersina has been allocated to a monotypic family by many
authors including Wetmore (1930, i960), Stresemann (1934), Hellmayr
75
(1936) and George (1962). Mayr & Amadon (195 1) recongized the family
Tersinidae "for the present," thus expressing uncertainty about the status
of the genus. Some authors have placed Tersina in a monotypic subfamily,
Tersininae (Mayr & Greenway 1956; Storer 1970) and others have depressed
it to a tribe, Tersinini (Beecher 1953), or to the level of a genus within the
tribe Thraupini (Sibley 1970).
The basis for recognition of a monotypic family, subfamily or tribe for
Tersina is that it is so different in its palatal structure from the typical tanagers
that it must be segregated from them. The counter argument emphasizes
the many similarities between Tersina and other tanagers and explains the
palatal and hyoideal differences as special adaptations associated with its
swallow-like method of capturing insects in flight. It should be noted that the
swallows (Hirundinidae) are one of the many groups of passerines which
have a cylindrical basihyale (George 1962:6). Thus convergence is a reason-
able explanation for the condition in Tersina. The tongue of Tersina was noted
by Lucas (1895b) as "hirundine in pattern" and the same type of tongue
"occurs among the swifts, and . . . probably ... in other insectivorous
birds ..." The similarities between Tersina and the typical tanagers involve
all characters other than those noted as differences. Lucas (1895b) pointed
out that the jaw, the nares and the ectethmoid are tanager-like. Lucas noted
that "the skull, in spite of its superficial resemblance to that of a swallow, is
structurally more nearly like that of such a typical tanager as Piranga erythro-
melas . . ." Beecher (1953:310) considered Tersina to be "anatomically close to
Calospi^a" [=Tangara] and Storer (1969) suggested that Tersina "may . . .
have been derived from a calliste-like ancestor." He further noted that "the
color and texture of the feathers are like those found in some callistes, and
the sexual dimorphism (blue males and green females) is like that in the Blue
Dacnis. The Swallow Tanager's bill is rather like that of a calliste but much
broader at the gape, a reflection of its fly-catching habits. Like the callistes,
Swallow Tanagers may also drift through the canopy of the forest in groups,
feeding on small fruits." The "callistes" are currently placed in Tangara
which is a synonym oiCalospi^a.
THE EGG WHITE PROTEIN EVIDENCE
In a study of the egg white proteins of passerine birds using starch gel
electrophoresis (Sibley 1970) I found that the pattern of Tersina was like that
of the tanagers and other members of the New World nine-primaried
assemblage. All of the members of this group were alike and it was not
possible to assess degrees of similarity and difference among them. Recently
the egg white proteins of Tersina have been compared with those of many
other passerines using the electrophoretic technique of isoelectric focusing
in acrylamide gel (abbreviated IFAG). The IFAG method has been described
by Sibley & Frelin (1972) and used in other studies by Sibley & Ahlquist
(1973) and Sibley (1973).
The greater resolving power of the IFAG technique permits detailed
comparisons of the isoelectric properties of homologous proteins in related
species. In the present study, using a pH 6-4 gradient, it was possible to
resolve at least 1 2 protein bands in the egg white of Tersina and other species
in the New World nine-primaried assemblage. The same basic pattern is
found in Tersina and in the more than 50 species of tanagers, wood warblers,
emberizine buntings and troupials that have been examined. Differences
among these species take the form of shifts in the isoelectric points and
76
relative amounts of homologous proteins. Homologies are easily determined
because the overall patterns are so similar.
After preliminary comparisons had shown that Tersina was a member of
the New World nine-primaried assemblage its egg white proteins were
compared in detail with those of the following species :
Tanagers (Thraupini): Thraupis episcopus, Thraupis sayaca, Buphonia lutei-
capilla, Piranga olivacea and Eucometis penicillata.
Wood- warblers (Parulini): Dendroica pensylvanica, Dendroica petechia,
Geothlypis trichas and Myioborus miniatus.
Honey-creepers (Coerebini): Coereba flaveola.
Buntings (Emberizini) : Emberi^a cirlus, Diuca diuca, Zonotrichia capensis and
Geospi^afortis.
Cardinal grosbeaks (Cardinalini) : Pheucticus ludovicianus, Passerina cyanea,
Saltator maximus.
Cardueline finches (Carduelini) : Acanthis flammea.
Troupials (Icterinae): Xanthopsar flavus.
The pH 6-4 IFAG pattern of Tersina egg white contains 12 protein bands.
All are identical in position and relative quantity to their homologs in
Thraupis episcopus, Thraupis sayaca, Piranga olivacea, Euphonia luteicapilla and
Coereba flaveola. The pattern oi Eucometis penicillata differs slightly from that of
these species in the positions of two bands. The patterns of the species of
Dendroica and Geothlypis are identical to one another but that of Myioborus
differs slightly in the position of one band. This difference makes Myioborus
virtually identical to Tersina and the genera which are identical to Tersina.
This suggests that these small differences are not consistent within recog-
nized groups and this is further indicated by most of the remaining species.
The patterns of the emberizine, cardinaline and icterine species are identical
or nearly identical to those of the tanagers, Tersina, etc. and I am not able to
find consistent differences correlated with currently recognized higher taxa.
The carduelines may differ enough to be separable from the emberizines etc.
but I cannot be certain that the observed differences will prove to be con-
sistent when many more species are compared.
What does seem clear is that closely related species do have identical
patterns. Within some genera there are small differences between species, but
it is far more common to find agreement between congeners to the finest
detail.
I interpret the evidence from all sources as indicating that Tersina is a
tanager and that its specialized characters are adaptations to its method of
feeding. There is no basis for taxonomic separation beyond the generic level
and the closest relatives of Tersina are probably Thraupis and Tangara.
DISCUSSION
Although the close relationship between Tersina and other tanagers may be
accepted by most avian systematists the more basic problem remains,
namely, the true generic alignments within the New World nine-primaried
oscines. I have discussed this question elsewhere (Sibley 1970: 106-108)
and Storer (1969) and Paynter & Storer (1970) also have commented upon it.
Briefly, the problem is as follows.
The groups herein recognized as forming the New World nine-primaried
assemblage (tanagers, wood-warblers, honey-creepers, buntings, cardinal
grosbeaks, troupials) are based primarily upon differences in the feeding
structures. For most of the groups bill shape is the most important defining
character. That all of these groups are closely related seems clear but whether
77
or not all "tanagers" are more closely related to other "tanagers" than to
some "wood-warblers" or "honey-creepers" is debatable.
Some steps toward a re-alignment of genera have been attempted. Beecher
(195 1) suggested that the Coerebidae is a composite of nectar-feeding
warblers and tanagers and proposed that Coereba, Ateleodacnis and Coni-
rostrum should be placed in the Parulidae and that the other "coerebid"
genera should be transferred to the Thraupidae. Skutch (1954:438; 1962)
and de Schauensee (1966:454) disagreed. Skutch (1962:113-116) argued that
Coereba is so distinct that it merits separation in a monotypic family and that
Conirostrum may be an emberizine. Lowery & Monroe (1968) adopted a
neutral position by placing Coereba and Conirostrum among a group of
"Genera Incertae Sedis" following the Parulidae.
Storer (1969) pointed out that the "pattern, color, and texture of plumage
offer many clues to relationships" and suggested that close relatives, in some
cases, have been placed in different "families". That this suggestion is correct
has been proved by the hybrids between the "coerebid" Cyanerpes cyaneus
and the "thraupid" Tangara nigrocincta reported by Delacour (1972a, b).
Delacour (1972a) also expressed his conviction that the "members of the
Sugar-bird family (Coerebidae) with the exception of those of the genus
Coereba, really are tanagers (Thraupidae)."
Although these debates have not solved any major problems they have
helped to define them and to make it clear that the present classification is
not a true reflection of natural relationships. The solution must be sought in
evidence which is unrelated to food habits because all characters associated
with feeding can be expected to reflect trophic adaptations and will thus
merely confirm the present arrangement rather than test its validity. The
procedure must involve a search for nearest relatives, mainly at the generic
level, without regard for the boundaries of currently recognized higher taxa.
The present study of Tersina is a step in this direction but many more difficult
questions remain unsolved. The IFAG patterns of the egg white proteins
seem likely to be helpful in identifying closely related taxa but material from
many more species must be obtained before such data will become significant.
AC KNOWLEDGMENTS
The specimens used in this study were made available by the generous
help of W. Belton, C. T. Collins, P. R. Davis, J. C. Doebeli, W. and E.
Fiala, H. Field, R. M. Fraga, S. Garber, J. Goodall, M. P. Harris, G. Hoy,
L. Kiff, R. W. Leeds, R. Leveque, E. S. Morton, S. Narosky, R. Pickering,
K. D. Pickford, H. Powell, N. Proctor, A. Ruschi, S. Rothstein, W. Spofford
and A. Williams.
I thank Jon E. Ahlquist, Elfriede Heaney, Dorothy J. Moore and Fred
C. Sibley for advice and assistance in the laboratory. The National Science
Foundation supported the study under grants GB-6192X and GB-36061.
References :
Beecher, W. J. 195 1. Convergence in the Coerebidae. Wilson Bull. 63: 274-287.
Beecher, W. J. 1953. A phylogeny of the oscines. Auk 70: 270-333.
Delacour, J. 1972a. Sugar-bird tanager hybrids. Avic. Mag. 78 : 48.
Delacour, J. 1972b. Hybrids sugar-bird X tanager. Avic. Mag. 78: 187-188.
George, W. 1962. The classification of the Olive Warbler, Peucedramus taeniatus. Amer.
Mus. Nov. 2103: 1-41.
Hellmayr, C. E. 1936. Catalogue of birds of the Americas. Field Mus. Nat. Hist., Zool. Ser.,
13, part 9 (Publ. No. 365): 458 pp.
Lowery, G. H., Jr. & B. L. Monroe, Jr. 1968. Family Parulidae, pp. 3-93 in Check-list of
Birds of the World, vol. 14, R. A. Paynter, Jr., ed. Cambridge, Mass.: Mus. Comp. Zool-
Lucas, F. A. 1895a. A new family of birds. Auk 12: 186.
78
Lucas, F. A. 1895b. Osteological and pterylographical characters of the Procniatidae.
Proc. U.S. Nat. Mus. 18: 505-507.
Mayr, E. & D. Amadon. 195 1. A classification of Recent birds. Artier. Mus. Nov. 1496: 1-42.
Mayr, E. & J. C. Greenway, Jr. 1956. Sequence of passerine families (Aves). Breviora (Mus.
Comp. Zool.) 58: 1— 1 1.
Paynter, R. A., Jr. & R. W. Storer. 1970. Introduction, pp. v-x in Check-list of Birds of the
World, vol. 13, R. A. Paynter, Jr., ed. Cambridge, Mass.: Mus. Comp. Zool.
Ridgway, R. 1895. Characters of a new American family of passerine birds. Proc. U.S.
Nat. Mus. 18: 449-450.
Ridgway, R. 1907. Birds of North and Middle America, pt. 4. U.S. Nat. Mus. Bull. 50:
973 PP-
Schaefer, E. 1953. Contribution to the life history of the swallow-tanager. Auk 70 : 403-460.
Schauensee, R. M. de 1966. The Species of Birds of South America. Livingston: Wynnewood,
Pa. 577 pp.
Sclater, P. L. 1862. Catalogue of a collection of American birds. N. Trubner & Co., London.
338 pp.
Sibley, C. G. 1970. A comparative study of the egg-white proteins of passerine birds.
Bull. Peabody Mus. Nat. Hist. 32: 1-131.
Sibley, C. G. 1973. The relationships of the silky flycatchers. Auk 90: 394-410.
Sibley, C. G. & J. E. Ahlquist. 1973. The relationships of the hoatzin. Auk 90: 1-13.
Sibley, C. G. & C. Frelin. 1972. The egg white protein evidence for ratite affinities. Ibis
114: 377-387-
Skutch, A. F. 1954. Life histories of Central American birds. Pac. Coast Avifauna 31 : 1-448.
Skutch, A. F. 1962. Life histories of honeycreepers. Condor 64: 92-116.
Storer, R. W. 1 969. What is a tanager ? Living Bird 8 : 1 27-1 36.
Storer, R. W. 1970. Subfamily Tersininae, pp. 408-409 in Check-list of Birds of the World,
vol. 13, R. A. Paynter, Jr., ed. Cambridge, Mass.: Mus. Comp. Zool.
Stresemann, E. 1934. Aves. In Handbuch der Zoologie, 7 Bd., 2 Ht., W. Ktikenthal & Th.
Krumbach, eds. Berlin: W. de Gruyter.
Vieillot, L. J. P. 18 19. Nouveau Dictionnaire d'Histoire Naturelle. Nouv. ed. Paris.
Wetmore, A. 1930. A systematic classification for the birds of the world. Proc. U.S. Nat.
Mus. 76: 1-8.
Wetmore, A. i960. A classification for the birds of the world. Smithsonian Misc. Coll.
139(11): 1-37.
On the taxonomic status of the genus Sauropatis
Cabanis & Heine (family Alcedinidae)
by A.K. Mukherjee & J. M. Dasgupta
Received 4th April, 1973
The genus Halcyon was described by Swainson (1821: text to pi. 27) with
Alcedo senegalensis Linnaeus as its type. His description of the genus is as
follows: — "Bill very long, straight, thick, the base broader than high; the
sides tetragonal: upper mandible very straight, the base rounded; under
mandible beneath carinated and recurved, the margins covered by those of
the upper. Nostrils basal, covered by a membrane, the aperture naked, linear
and oblique. Tail mostly moderate. Feet gressorial: interior fore-toe small
or wanting."
Cabanis & Heine (i860) erected the genus Sauropatis with Halcyon sanctus
Vigors & Horsfield as its type-species. This species was distinguished from
Halcyon senegalensis (Linnaeus) by its short and broad bill, the lower mandible
being much more curved upwards and the bill being black and not red.
Sharpe (1868) pointed out that the genus Halcyon had earlier been divided
into many genera by different workers, but those differences were merely
those of "Style of plumage and not those of forms". He, therefore, synony-
mized Sauropatis with Halycon which is represented in the Ethiopian, Indian
and Australian Regions and parts of the Palaearctic Region. In this genus, he
included as many as 3 6 species of which four belonged to the Indian Region,
79
namely, Halcyon coromanda, Halcyon smyrnensis, Halcyon pileata and Halcyon
chloris.
Blanford (1895: 135-137), Mathews (1912: 108-109), Oberholser (1919)
and Baker (1927: 274-278) have advocated the segregation of Sauropatis
Cabanis & Heine from Halcyon Swainson, while Sharpe (1870), Biswas
(1953: 32), Smythies (1953: 345-352), Peters (1945: 207-209) and Ripley
(1961: 220-221) have synonymized Sauropatis with Halcyon.
Except Sharpe (1868), however, none of the above-mentioned authors has
assigned any reason for his action.
From an examination of type-species of the genera Halcyon and Sauropatis ',
we find the following differences :
Halcyon Sauropatis
1. Bill rather large, broad at base; Bill short, broad; lower man-
culmen straight, rounded above. dible much curved upwards.
2. Bill generally red. Bill all black or nearly so.
3. 1 st (outermost) primary half the 1st (outermost) primary sub-
length of the 2nd; 3rd primary equal to the 2nd; 3rd primary
being the longest. almost equal to the 2nd.
4. 1 st (outermost) primary shorter 1st (outermost) primary longer
than 5 th and 6th primaries. than 5 th and 6th primaries.
5. A white wing-patch invariably White wing-patch absent,
present on the bases of the pri-
maries, except in Halcyon coro-
manda.
6. Inhabits inland pools, fresh- Inhabits mostly coastal regions
water rivers and estuaries. and sometimes estuaries.
7. Nests are made in tunnels on Nests are made generally in
river banks, and sometimes in termite mounds or in clay nests
holes of tree-trunks near rivers. of ants.
8. Clutch-size varies from 2 to 6, Clutch-size generally 3.
but generally more than 3.
AH & Ripley (1970: 95-99), while giving the description of the genus
Halcyon, remarked that "In one species chloris, lower mandible is much more
curved upwards". We find that this character is not only confined to chloris
but also to another species, namely, sancta. These species are not only
distinguishable from other species of Halcyon on morphological characters
but they also have different habits, such as in nesting, habitat preferences,
feeding, etc. Hence, there is ample justification to group them {chloris, sancta)
in a genus separate from other representatives of Halcyon. We would,
therefore, resuscitate the genus Sauropatis Cabanis & Heine to accommodate
these two species.
The genus Sauropatis is more or less limited to the humid tropical coasts
of south-eastern India, Malaysia, Polynesia and Australasia.
We are indebted to the authorities of the American Museum of Natural
History, New York, for the loan of some specimens for our study. We are
thankful to the Director, Zoological Survey of India, for extending facilities
for the present work, and to Dr. B. Biswas, for critically reading through the
manuscript and suggesting improvements.
References:
Ali, S. & Ripley, S. D. 1970. Handbook of the birds of India and Pakistan. 4. Bombay: Oxford
University Press.
Baker, E. C. S. 1927. Fauna of British India, birds. 4. London: Taylor & Francis.
Biswas, B. 1953. A check-list of genera of Indian birds. Rec. Indian Mus. 50.
80
Blandford, W. T. 1895. Fauna of British India, birds. 3. London: Taylor & Francis.
Cabanis, J. & Heine, F. i860. Museum Heineanum. 2. Berlin.
Mathews, G. M. 191 2. New generic names for Australian birds. Austr. Avian Rec. 1.
Oberholser, H. C. 191 9. Revision of the subspecies of the White-collared Kingfisher,
Sauropatis Moris (Boddaert). Proc. U. S. Nat. Mus. 55: 351-354.
Peters, J. L. 1945. Check-list of birds of the world. 5. Cambridge, Mass.: Harvard Univ. Press.
Ripley, S. D. 1961. A synopsis of the birds of India and Pakistan. Bombay: Bombay Nat. Hist.
Soc.
Sharpe, R. B. 1868-71. A monograph of the Alcedinidae. London: the author.
Smythies, B. E. 1953. The birds of Burma. 2nd ed. Edinburgh: Oliver & Boyd.
Swainson, W. 1821. Zoological illustrations. 1. London: R. & A. Taylor.
Ortolan and Blue Rock Thrush in Sierra Leone
by G. D. Field
Received 6th February, 197}
Few records of wintering Ortolans Emberi^a hortulana are available from
West Africa and Moreau (1972) only notes "a few . . . found wintering in
Nigeria south to about 9 50' N." and a double passage on the lower Senegal
River. In the first week of January 1973 on an expedition to the Loma
Mountains in N.E. Sierra Leone, 9 13' N. and n° 06' W., I found the species
plentiful on the steep south-east face of Bintimane, the highest point of the
mountains. On this side the gallery forest ends at approximately 1,000 ft.
below the summit, 6,390 ft., and the 600 or so feet above the forest consists
partly of precipitous slabs of bare rock, partly of steep grass and herb
covered slopes, interspersed with large boulders and a few scattered trees.
By January the whole slope had been burned except for some moister folds
where tiny springs still flowed. The buntings could be found over the whole
of this slope but not on the gentler bare ground above leading to the summit.
They were in company with the pipits, Anthus trivialis and A. similis, both
very common, and to a lesser extent with the native bunting Emben\a
tahapisi, spending most of their time on the ground but perching freely on
the burnt herb stems and occasionally in trees. Plumage was very variable,
ranging from bright, clear yellow and cinnamon-rufous underparts to drab
whitish with pale pinkish bellies, while some had rather heavily streaked
breasts. They were present throughout the five days of my stay and were not
seen outside the one area. Though I once had six together on or beside one
small burnt bush they did not normally flock closely and, assuming an even
spread over the slope, I should tentatively put the population somewhere
about fifty individuals.
Until recently records of the Blue Rock Thrush Monticola solitarius were
almost equally scanty for West Africa south of the Sahara, but it has now
been reported from Gambia (McGregor & Thomson 1965), northern and
southern Nigeria, Senegal and Mt. Nimba in Liberia (Moreau 1972). In
early December 1965 I noted at least three individuals on the summit slopes,
above 5,500 ft., of the Tingi Mountains, thirty miles south-west of the Lomas
and very similar in formation and vegetation. At the time I was not sure
whether this was a genuine wintering ground or merely a freak occurrence.
However, the original sighting is confirmed by its presence in the Lomas on
the same slope as the Ortolans and right up to the summit. Birds were seen on
several occasions and five individuals noted on one morning, making it here
more common than M. saxatilis, though the latter also occurs on the plateau
below the summit slope and along the edges of the galleries. Where the two
occurred together M. saxatilis was once noted chasing M. solitarius from a
favourite boulder. Taken together, these two records suggest that M.
solitarius regularly winters in Sierra Leone in small numbers in the available
areas above 5,000 ft.
References :
McGregor, I. A. & Thomson, A. L. 1965. Blue Rock-thrush Monticola solitarius in the
Gambia. Ibis 107: 401.
Moreau, R. E. 1972. The Pa/aearctic- African Bird Migration Systems. London & New York:
Academic Press.
The migratory movements of the Pygmy Kingfisher
Ceyx picta natalensis in the Republic of Zaire
by A. Prigogine
Received 26th February, 1973
It is some years since various authors, especially Benson (1964: 57-60), have
shown that the southern form of the Pygmy Kingfisher Ceyx picta (Boddaert),
namely C. p. natalensis (Smith), has migratory movements, while recently
Clancey (1972) has drawn attention to some off-season records of this form
from the Bwamba Forest, extreme western Uganda. Moreau (1966: 243) cites
C. picta among intra- African migrants. On the other hand, it is curious that
Mackworth-Praed & Grant (1970: 424) make no mention of any movement
in C. p. natalensis. With few exceptions natalensis occurs in Zambia, Malawi
and Rhodesia, and in territories to the southward, from September to March
(Benson 1964: 58). Breeding takes place in Zambia, Malawi and Rhodesia in
the same period, with a peak of egg-laying in November (Benson 1963 : 632;
Benson et al. 1964: 59). Benson (1964: 59) also demonstrated that these
southern populations of natalensis migrate northward, particularly to north-
eastern Zaire (Kivu), Rwanda, Uganda and Kenya (see also Clancey 1972,
who makes no mention of Benson's data). The northernmost record which
Benson gives is from Lake Albert, between Hoima and Bugoma, since when
I have even found it at Paulis (Isiro), in the Upper Uele at 2 5 3 ' N., 27 5 8 ' E.
If the status of natalensis is well established in Zambia and in other countries
to the southward, the situation is on the other hand much less clear in Zaire.
This is because in certain parts of Zaire natalensis appears only in a deter-
minate period, to occur at this time alongside a population of C. p. picta
which is resident throughout the year. So the presence of natalensis can
remain unperceived, and as I have already noted myself many specimens
of natalensis have been confused with picta or vice versa.
In the first place, it must be remembered that in the southern part of Zaire
there exists a population of natalensis which seems to have exactly the same
rhythm as that in Zambia (Benson et al. 1971: 156). Bohm had already
collected this form in the Katanga, near the Rivers Dikulwe and Lufira, but
his specimens were recorded by Schalow (1886: 421, 432) and Matschie
(1887: 151) merely as Ispidina picta. Chapin (1939: 286) has cited natalensis
from the Katanga, but was unable to examine personally any specimens
from this part of Zaire, while the record by Neave (1910: 109) listed is in
reality from Zambia. Subsequently Verheyen (1947: 4) has recorded
natalensis from Musosa (collector H. J. Bredo), without giving the date of
collection; actually the specimen is a female collected some time in the
period September/November. Schouteden (195 1: 30-32) has given all the
localities in Zaire, Rwanda and Burundi in which the form natalensis has been
collected, and has mentioned in addition Albertville, in the northern Katanga.
82
Verheyen (1953: 347) was the first to note dates of collection, of five
specimens from the Upemba National Park. They fall within the period late
August to mid-April (the specimen shown as dated 27th November is
actually labelled 27th August). It seemed to Verheyen that "Pespece est
sujette a des migrations (locales?)". He estimated that breeding was in
January/February, according to the age of birds examined and the state of
their plumage. Later, Schouteden (1965 : 37) cited all the specimens emanating
from Dilolo and Kolwezi under the name Ispidina picta, and recorded their
dates of collection as falling between mid-August and the beginning of
February. Finally, Schouteden (1971: 92) summarised all the specimens
collected in the Katanga as a whole as coming within the period 9th August
and the end of March. So, in summary, in the Katanga the situation for
natalensis corresponds well with that described for Zambia. The birds arrive
in August, breed and then disappear, the latest most often in March, as we
shall see in detail below.
What happens to the natalensis of this population in the ensuing period ?
It is not possible to distinguish these birds from those of other populations
nesting to the southward, and they must be considered together. The
information in the literature concerning natalensis collected outside the
breeding range is extremely fragmentary. The first to have recorded natalensis
only a little to the south of the equator is without doubt Gyldenstolpe (1924:
276), who mentions a female collected near Lake Chahafi, Uganda. Chapin
(1939: 286) in turn noted that in the museum at Tervuren there were several
natalensis from Kisenyi, the vicinity of Lake Kivu and the Rutshuru Plain.
Schouteden (1942: 334) recorded the collection of a female of natalensis at
Kirinda, Rwanda on 25 th August, although personally I attribute it to the
nominate form. Schouteden (195 1 : 30-32) recorded several further specimens
from Kivu, at Kalembelembe, and from Rwanda/Burundi, at Kisenyi and
Rugegera, but did not give collecting dates. It can also be noted that
Aurelien (1957: 206), in describing Ispidina picta from Rwanda, writes that
"les auriculaires portent une tache bleue". This description corresponds
with natalensis, which would accordingly appear to be common enough in
Rwanda and Burundi (actually Aurelien sent only two specimens of natalensis
to Tervuren).
Some specimens from Zaire, Rwanda and Uganda have been recorded by
Benson (1964: 59, 66), in particular from Kivu, at Lwiro; from Rwanda, on
the River Kagera, and at Gisagara and Kisenyi; and from extreme western
Uganda, in the Bwamba Forest (and see also Clancey 1972). Subsequently to
Benson, Schouteden has given a list of specimens of Ispidina picta in the
collections at Tervuren, from Rwanda (1966a: 57), Burundi (1966b: 39) and
Kivu (1968 : 120), but did not separate the forms picta and natalensis.
For this reason, in order to clarify the occurrence of natalensis in Zaire and
adjacent territories to the east, I have examined all the specimens of C. picta,
totalling more than 600, in the collections of the Musee royal de l'Afrique
Centrale, Tervuren and in the Institut royal des Sciences Naturelles, Bruxel-
les. I have also taken note of a certain number of specimens recorded in the
literature, in each such case expressly indicating this. In all, exclusive of the
10 recorded by Clancey (1972) from the Bwamba Forest, I have been able to
bring to light 91 specimens of natalensis, six of them without date of collec-
tion. Eighty-five of them appear in Table 1, which shows their monthly
distribution for the different regions considered. The Map indicates the
position of the localities where natalensis has been collected, and the
83
corresponding season of the year. Finally, in Table 2, I have drawn up a list
of all the specimens of natalensis which form the basis of this study.
TABLE 1
Distribution of specimens of Ceyx picta natalensis by months (exclusive of ten recorded by
Clancey 1972 from Bwamba, Uganda between 25th April and 19th July).
J F MAMJJIASOND Total
Southern Katanga (a) 3 6 421 2 8 10 6 5 47
Northern Katanga 11 2
Kasai 1 1 1 3
Lake Leopold II 11
Lower Congo 1 1
Maniema 1 1 1 3
Southern Kivu (b) 1 1 1 3
Burundi 1 (c) 1
Rwanda (b) 22 i(d) 5
Idjwi Island 24 6
Northern Kivu &
Uganda (b) i(e) 2 3 2 3 i(f) 12
Upper Uele 1 (g) 1
TOTAL (without
southern Katanga) 1 24739101 1 38
(a) Includes two specimens collected by Bohm.
(b) Includes specimens cited by Benson (1964: 59) from Lwiro, Kagera and Bwamba.
(c) Collected at Bujumbura 1st March, wing 55 mm; a second specimen from
Bujumbura, undated, has wing 50-5 mm only, but there is a well developed patch
of blue on the ear-coverts.
(d) Collected at Kisenyi in Nov., wing 55-5 mm; but in poor condition, blue on ear-
coverts little visible.
(e) Collected near Lake Chahafi 6th Feb.; wing 54 mm.
(f) Collected by Grauer at Beni in Sept. (Sassi 1912: 365).
(g) Collected at Paulis (Isiro) 19th March; wing 55 mm.
TABLE 2
List of specimens examined or cited from Zaire, Rwanda, Burundi and western Uganda (a)
A. Southern Katanga: 1 Bunkeya 20. II; 1 Buvumba (river) 23. IX; [1 Dikulwe (river)
25.XI]; 1 Kansenia 11. Ill; 25 Kasaji 16.VIII/9.II; 2 Kasapa V, IX; 1 Kiambi 5.XII;
2 Kipopo 3 1. Ill, 28.IV; 1 Kiswishi 15.X; 2 Kolwezi 20.III, 27. XII; [1 Lufira (river)
10.X]; 1 Lupando 14.IX; 1 Mpala 19. X; 1 Musosa IX-XI; 1 Mweru (lake) 23.I; 1
Tanganyika (lake) undated (b); 5 Upemba Nat. Park 27.VIII/15.IV.
B. Outside southern Katanga:
Northern Katanga: 2 Albertville 24.IV, 9.VIII.
Kasai: 1 Gandajika 16. IV; 1 Kasanza 2.V; 1 Luluabourg 2.VII.
Lake Leopold II: 1 Bokalakela 2 5. VIII.
Lower Congo: 1 Kinshasa 4.V.
Maniema: 1 Kalembelembe undated; 2 Kampene 2.VII, 14.VIII; 1 Kasongo 8.IV.
Southern Kivu: 1 Hombo 25. V; 1 Kamituga 28.IV; 1 Lwiro 4. VII (c).
Burundi: 2 Bujumbura 1 .III, undated.
Rwanda: 2 Astrida undated; 1 Gisagara 7.V; 1 Kagera 27.VII (c); 3 Kisenyi VII, XI,
undated; 1 Rugegera 5.V.
Idjwi Island: 6 Idjwi 9.VII/24.VIII.
Northern Kivu: 1 Beni IX; 1 Butembo 15. VI; 1 Etaetu 21.V; 1 Etembo VI; 2 Mutsora
1 1. VIII, 27.VIII; 1 Mutwanga 22.V; 1 Rutshuru (plain) VI.
Uganda {near frontier with Zaire) (d): 1 Bwamba (forest) 3. VIII; [2 Bwamba (forest)
n.VII, 20.VII]; 1 Chahafi (lake) 6.II.
Upper Uele: 1 Paulis (Isiro) 19. III.
(a) Specimens cited, not examined by myself, are shown in brackets. Not included are
10 specimens from the Bwamba Forest, western Uganda, collected between 25 th
April and 19th July (Clancey 1972).
(b) Specimen collected in 1883 by Storms.
(c) Examined by Mrs. R. T. Chapin (previously cited by Benson 1964: 59).
(d) See also second sentence under (a).
84
Distribution of specimens of Ceyx picta natalensis by months.
If we consider in the first instance the southern Katanga, 42 out of 47
specimens were collected between September and March. There are two for
August (16th August, Kasaji; 27th August, Upemba National Park). On
the other hand, two were still present in April (1 5th April, Upemba National
Park; 28th April, Kipopo), and one even in May (Kasapa, near Lubumbashi,
without indication of day of collection). So it can be admitted that natalensis
is present from September to March, but that some individuals arrive as
early as mid-August and the last leave in April/May. The situation thus
agrees well with that shown by Benson (1964: 5 8) for Zambia, etc.
Apart from the southern Katanga, I have been able to take count of a total
of 38 specimens of which 33 fall within the period April/August, exactly the
period when natalensis is almost completely absent from the southern
Katanga. Furthermore, these figures are exclusive of ten specimens of
natalensis mist-netted in the Bwamba Forest, Uganda between 25th April
and 19th July, reported on by Clancey (1972), for which the exact dates
within this period are not indicated.
In the northern Katanga, the dates of two specimens collected at Albert-
ville seem to correspond respectively with northward and southward pas-
sages — 24th April and 9th August. There are records of natalensis from the
Kasai and even along the River Congo in the districts of the Lower Congo
and Lake Leopold II.
But it is above all in the eastern part of Zaire and adjacent Uganda that
natalensis seems to make the longest movements as already indicated by
Benson (1964: 59) and recently by Clancey (1972). In fact natalensis has been
collected in Maniema, in different parts of Kivu, on Idjwi Island (specimens
from this locality had previously been confused with nominate picta in
Prigogine 1967: 260), in Rwanda and Burundi, just across the Zaire border
in the Bwamba Forest, Uganda, and in the Upper Uele.
The female collected on 6th February near Lake Chahafi, Uganda and the
male on 1st March at Bujumbura, Burundi represent exceptionally advanced
dates. On the other hand, the latest specimens of natalensis have been
collected at the end of August (24th August on Idjwi Island; 27th August at
Mutsora, northern Kivu). But an isolated specimen has been collected in
November, without indication of the day of collection, at Kisenyi, Rwanda.
It is not possible to say, in the case of specimens collected outside the period
April/ August, whether it is a question of migrants, or of natalensis remaining
in their winter quarters.
In the migratory period, natalensis does not seem to have a preference for
any particular biotope. On the one hand, specimens have been collected in
open habitats as for example at Bujumbura, Kisenyi and on the Rutshuru
Plain. On the other, they have also been found in heavy forest as at Bwamba,
Hombo, Etaetu and Etembo.
The stay of natalensis in its winter quarters corresponds with sexual
inactivity. Let us examine, then, the breeding period of nominate picta.
According to Chapin (1939: 286) nesting near the equator may continue
throughout the year, but further to the north it probably takes place from
March to May or June, while to the south Chapin expected it to commence
at the start of the rains (that is to say, in September/October). According to
Traylor (1963 : 92), the female and three eggs collected in March at Calulo,
in Angola at io° S. (Lynes & Sclater 1934: 38), belong to nominate picta.
Around Kamituga, at ca. 3 S., I found a nest with three eggs on 25th
October (Prigogine 1961:253) and another with four eggs on 29th August
(Prigogine 1972: 207). Based on all my records from this region, including
also those of young, I conclude that breeding is from August to October and
from January to March, in fact probably the whole period mid-August to
March (Prigogine 1971 : 70). It would seem that, when natalensis is present in
this region, from April to August, and is found alongside the nominate form,
there is no reproductive activity, and this would favour the segregation of the
two forms. But, as we shall see, this does not seem to be the case everywhere,
particularly in the southern part of Zaire and further southward.
In principal, the integration between natalensis and picta could take place in
regions where natalensis breeds but where there exists also a resident popula-
tion of nominate picta nesting at the same season, as in Angola commencing
at the start of the rains (Traylor, 1963 : 92, also gives a record of a juvenile
nominate picta from Dundo for as early as August). White (1965: 226)
records "some integration" in the Kasai, but in the collection at Tervuren
there is no specimen from the Kasai showing characters intermediate between
natalensis and nominate picta.
86
On the other hand, Benson (1964: 60) records a male from Vila Texeira de
Sousa, Angola which seems intermediate. In general, such specimens could
occur in a belt, where natalensis and picta could be in contact, during the
breeding season of natalensis and which would traverse Africa in the vicinity
of ii° S. in Angola and 5 S. in Tanzania. It is difficult to trace such a band
across Zaire. According to the specimens examined, the separation between
natalensis and picta could be along a line which passes close to Albertville
and a little to the north of Kasaji. But on the other hand Benson (1964: 57)
records two specimens of picta from Salujinga in the extreme north-west of
Zambia, thus to the south of Kasaji. Indeed, Benson & Irwin (1965: 3)
record a further two specimens of this form from Salujinga. The situation
in this part of Africa, then, seems complicated, and one must await some
intermediates. Such could "behave" either like natalensis or like picta. In the
first case, they would have seasonal migrations; in the second, they would
be resident throughout the year. Two females in the Institut royal des
Sciences Naturelles could be intermediate. Both were collected at Mutsora,
in the vicinity of Ruwenzori, on 16th April and 1 ith August (as they cannot
definitely be attributed to natalensis, they are not included in the tables). The
first one has a mauve spot washed with blue on the ear-coverts; the second
shows the start of a blue spot. Nevertheless the black of the crown is more
intense in both specimens than in specimens of true natalensis.
Furthermore, in the collection at Tervuren I have found six specimens
which could also be intermediates. Although the blue spot on the ear-
coverts is not visible — perhaps partly due to bad preparation — the colour of
the crown is like that of natalensis. In two of them, from Bolafa (a little to the
south of Bumba, in the Equator province), without indication of date of
collecting, the underside is also a relatively pale orange-brown, as in natalensis.
In two others, collected in April and September at Buta, Lower Uele the
crown is as in natalensis, but the underside is darker, as in nominate picta.
Finally, in two from Etaetu, northern Kivu, the crown is more blackish, and
they seem intermediate between natalensis and picta. As to the colour of the
underside, it is pale in one of these two specimens, collected in January, but
darker in the other, collected in June. The wing-length varies in these six
specimens as a whole from 50 to 52-5, mean 51 mm, these figures agreeing
with the nominate form. In summary, their status is dubious. Perhaps they
are intermediate. Account cannot be formally taken of them in this study, but
it is worth drawing attention to them.
It may be noted that 3 5 specimens of natalensis from further north than the
southern Katanga range in wing-length from 50 to 56, mean 54-0 mm, which
is in close agreement with the figure 54-3 mm given by Benson (1964: 57)
for 166 specimens.
Summary: (1) In order to clarify the presence of Cejx picta natalensis in
Zaire and adjacent country to the east, more than 600 specimens of C. picta
in the collections of the Musee royal de l'Afrique Centrale (Tervuren) and
Institut royal des Sciences Naturelles (Bruxelles) were examined.
(2) It has been possible to discriminate 91 specimens of natalensis, six of
them without date of collection. Account is taken in this figure of a certain
number of specimens previously recorded in the literature (but not of 10
specimens recently recorded from the Bwamba Forest by Clancey).
(3) In the southern Katanga, 42 out of 47 specimens of natalensis were
collected between September and May. This corresponds well with the period
in which this form is present in Zambia and further south.
87
(4) In regions to the northward, 33 out of 38 specimens of natalensis fall
in the period April to August, that is to say, the period in which natalensis is
absent from the southern Katanga.
(5) The segregation between natalensis and nominate picta is favoured by
the fact that the reproduction of the latter does not take place during the
period corresponding to the presence of natalensis in its winter quarters. On
the other hand, in the southern part of Zaire and to the southward, where
there exists a resident population of nominate picta, the two forms nest at the
same time, and non-integration is possible. But attention is drawn to several
specimens which could be intermediate.
Acknowledgements: I am most grateful to Mr. C. W. Benson, who has
provided me with much valuable information and who has had the extreme
kindness to translate my paper.
I am also much obliged to Mrs. R. T. Chapin, who examined the specimens
in the American Museum of Natural History; and to Dr. H. Schifter (Vienna)
and Mr. Devillers (Brussels), who were very helpful during my examination
of the material under their care.
References:
Aurelien, Frere. 1957. Les oiseaux du Ruanda-Urundi. 1. Astrida, Rwanda.
Benson, C. W. 1963. Breeding seasons of birds in the Rhodesias and Nyasaland. Proc.
i$th Internat. Orn. Congr. II: 623-639.
— 1964. Some intra- African migratory birds. Puku 2 : 5 3-66.
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. Stuart. 1971. The birds of
Zambia. London: Collins.
Benson, C. W., Brooke, R. K. & Vernon, C. J. 1964. Bird breeding data from the Rhode-
sias and Nyasaland. Occ. Pap. Nat. Mus. S. Rhod. 27B: 30-105.
Benson, C. W. & Irwin, M. P. Stuart. 1965. Some birds from the North-Western Province,
Zambia. Arnoldia (Rhodesia) 1 (29): 11 pp.
Chapin, J. P. 1939. The birds of the Belgian Congo: Part 3. Bull. Amer. Mus. Nat. Hist. 75.
Clancey, P. A. 1972. Miscellaneous taxonomic notes on African birds. Durban Mus. Novit.
9(11): 145-146.
Gyldenstolpe, N. 1924. Zoological results of the Swedish expedition to Central Africa
1921. Vertebrata. I. Birds. Kungl. Sv. Vetensk. Akad. Handl. 3 (1), 3: 1-326.
Lynes, H. & Sclater, W. L. 1934. Lynes- Vincent tour in Central and West Africa in 1930-
31. Part II. Ibis (13) 4: 1-5 1.
Mackworth-Praed, C. W. & Grant, C. H. B. 1970. African handbook of birds. Series III,
Vol. 1. London: Longmans.
Matschie, P. 1887. Der ornithologische Nachlass Dr. Richard Bohm's. II. Journ. Ornith.
35: 135-156.
Moreau, R. E. 1 966. The bird faunas of Africa and its islands. London : Academic Press.
Neave, S. A. 1910. On the birds of Northern Rhodesia and the Katanga District of Congo-
land (part 1). Ibis (9) 4: 78-155.
Prigogine, A. 1961. Nids et oeufs recoltes au Kivu (Republique du Congo). Rev. Zool. Bot.
Afr. 64: 248-264.
— 1967. La faune ornithologique de Tile Idjwi. Rev. Zool. Bot. Afr. 75 : 249-274.
— 1971. Les oiseaux de l'ltombwe et de son hinterland. Ann. Mus. Afr. Centr. 8, Sc.
Zool. 185.
— 1972. Nids et oeufs recoltes au Kivu. 2. (Republique du Zaire). Rev. Zool. Bot. Afr. 85 :
203-226.
Sassi, M. 191 2. Beitrag zur Ornis Zentralafrikas. Ann. k. k. Naturhist. Hofmus. 26: 347—393.
Schalow, H. 1886. Der ornithologische Nachlass Dr. Richard Bohm's. Journ. Ornith. 34:
409-436.
Schouteden, H. 1942. Contributions a la faune ornithologique du Ruanda-Urundi. I.
Collection J. A. Lestrade. Rev. Zool. Bot. Afr. 36: 329-342.
— 195 1. De Vogels van Belgisch Congo en van Ruanda-Urundi. IV. Ann. Mus. Congo,
C. Zool. (4) 3, fasc. 1.
— 1965. La faune ornithologique des territoires de Dilolo et Kolwezi de la Province du
Katanga. Mus. Roy. Afr. Centr. Doc. Zool. 9.
— 1966a. La faune ornithologique du Rwanda. Mus. Roy. Afr. Centr. Doc. Zool. 10.
— 1966b. La faune ornithologique du Burundi. Mus. Roy. Afr. Centr. Doc. Zool. 11.
88
Schouteden, H. 1968. La faune ornithologique du Kivu. I. Non passereaux. Mus. Roy.
Afr. Centr. Doc. Zool. 12.
— 1 97 1. La faune ornithologique de la Province du Katanga. Mus. Roy. Afr. Centr. Doc.
Zool. 17.
Traylor, M. A. 1963. Checklist of Angolan birds. Pub/. Cult. Comp. Cia. Diamant. Angola 61.
Verheyen, R. 1947. Notes sur la faune ornithologique de l'Afrique Centrale. VIII. Bull.
Mus. Hist. nat. Beige 23 (10): 4 pp.
— 1953. Exploration du Pare National de VUpemba. Fasc. 19. Oiseaux. Bruxelles: Inst.
Pares Nat. du Congo Beige.
White, C. M. N. 1965. A revised check list of African non-passerine birds. Lusaka: Government
Printer.
Sped
rimen data on Bucorvus leadbeateri
by R.K. Brooke & A. C. Kemp
Received 1st March, 197}
Verheyen (1953) appears to be the only author to discuss moult in the
Southern Ground Hornbill Bucorvus leadbeateri (Vigors), although Stresemann
& Stresemann (1966) have discussed it in the related Abyssinian Ground
Hornbill B. abyssinicus (Boddaert). R. K. B. had the opportunity to examine
while fresh a dying subadult female B. leadbeateri brought into the National
Museum in Bulawayo (NMB) on 6 June, the existing material preserved
there, in the Queen Victoria Museum in Salisbury (QVM) and the Durban
Museum (DM). A. C. K. examined the material preserved in the Transvaal
Museum in Pretoria (TMP).
Details of the state of moult and wear of the primaries, secondaries and
rectrices are shown in Table 1. It will be noted that only three specimens
show active primary moult (code letter G in Table 1) and these were col-
lected in January, March and June. The bird taken on 3 October, probably
about two years old since the egg-laying season in Rhodesia is August to
November, shows a simple symmetrical descending mode of replacement
of the primaries. However, a bird taken on 23 October, though long out of
the nest, shows no sign of moulting, even though the adult female, collected
on the following day in the same area, had an egg in the upper part of the
oviduct. The 23 October bird is probably about one year old. In the 14
adults and subadults moult is diagonal or asymmetrical starting in one wing
after the other, although in which wing is a matter of individual variation.
The data in Table 1 suggest that in B. leadbeateri the same pattern, with
individual variation, of primary moult occurs as in B. abyssinicus studied by
Stresemann & Stresemann (1966), i.e. diagonal stepwise descending moult
in primaries 1 to 7 and diagonal stepwise ascending moult in primaries 10 to
8.
Active secondary moult was evident in three specimens, the two from
June and January which also showed primary moult, and a third which
showed no primary moult or data on collection. It may well be that more
specimens than the three noted showed active moult of the secondaries but
this aspect was not checked by R.K.B. The fresh B. leadbeateri had 17
secondaries and the data in Table 1 suggest that asymmetry is less marked in
moult of the secondaries than in the primaries, and also that there are three
centres at which moult starts, i.e. secondaries 1 (the outermost), 10 or
thereabouts and 16. There are four alula quills and their mode of moult is a
descending one. The fresh B. leadbeateri had a subalular apterium similar to
that described for B. abyssinicus by George & Casler (1972).
Active rectrix moult was present in specimens taken in November,
89
TABLE i
Details of moult in the primaries, secondaries and rectrices of Rucorvus leadbeateri
A. PRIMARIES
Material
NMB
Age & Sex
9
9
9
6*
Subad.
Ad.
Ad.
Ad.
Ad.
Juv.
Ad. not
sexed
Ad.
Ad.
Ad.
Ad.
Ad.
Date
6 June
12 Aug.
7 Sept.
4 Nov.
14 Dec.
3 Oct.
3 July
Aug.
8 July
no date
21 Dec.
23 Jan.
24 Oct.
QVM
DM
Ad.
(on point of laying)
Juv. 6* 23 Oct.
Ad. 6* 25 March
Subad. ? 29 July
wing
Pr
manes
1
2
3
4
5
6
7
8
9
10
L
W
\\"
\V
N
F
G
O
F
F
F
R
W
w
W
W
N
F
G
O
F
F
L
N
N
F
F
W
W
W
F
N
N
R
F
F
W
w
w
W
N
F
N
N
L
N
N
F
o
o
F
O
o
F
O
L
F
F
F
F
o
F
w
w
F
w
R
N
W
N
N
N
O
o
\v
N
N
L
N
O
W
O
o
O
o
o
o
W
R
N
w
o
\v
o
O
o
N
o
N
L
F
F
o
o
o
O
o
o
o
O
R
F
F
o
o
o
o
o
o
o
O
L
O
N
N
N
F
w
w
w
F
N
R
O
O
N
N
F
w
w
w
F
N
L
N
N
F
W
W
N
F
w
N
N
R
N
F
F
w
\V
\V
\v
N
F
N
L
N
F
F
\v
N
N
F
o
N
N
R
N
N
F
F
W
W
W
N
F
N
L
N
w
W
W
w
w
N
F
\V
N
R
N
N
N
N
w
w
N
N
N
N
L
N
N
N
N
N
w
N
W
N
N
R
N
N
N
N
\V
N
N
w
N
N
L
F
G
W
N
W
G
W
w
N
N
R
N
F
G
W
W
N
G
\v
N
G
L
N
F
F
W
N
N
N
F
W
N
R
N
F
F
w
\V
\V
N
F
\v
F
L
N
N
N
N
N
N
N
N
N
N
R
N
N
N
N
N
N
N
N
N
N
L
F
F
N
F
o
G
o
O
O
—
R
F
F
N
N
F
O
o
O
'F
_
R
N
N
N
N
W
F
F
o
N
F
B. SECONDARIES
Material Age e> Sex Date
NMB
Subad. $
6 June
Wing
TMP
d
f
Ad.
Ad.
<5
no date \
23 Jan. /
Some :
C. RECTRICES
Material
Age <&■ Sex
Date
Tail
R
NMB
a
Subad.
9
6 June
b
Ad.
9
12 Aug.
c
Ad.
9
7 Sept.
d
Ad.
<J
4 Nov.
e
Ad.
9
14 Dec.
f
Juv.
o"
3 Oct.
QVM
Ad.
0*
25 Mar
TMP
b
Ad.
9?
Aug.
c
Ad.
o*
8 July
d
Ad
9
no date
e
Ad.
9?
21 Dec.
f
Ad.
6"
23 Jan.
g
Ad.
9
24 Oct.
(on point
of laying)
h
juv.
S
23 Oct
DM
Subad
+
29 July
Code:
D =
dropped
F
= fresh
G = growing
R =
right
W
= worn
— = damaged
Secondaries
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
F GOOOOF GF F F WWWWF O
FFOODFOGFFFNNNNFO
Some feathers growing in secondaries of each wing
5
4
3
2
1
1
2
3
4
5
W
F
N
O
N
N
O
N
F
W
F
W
O
N
O
N
o
F
W
N
O
N
W
\V
O
W
o
N
w
F
F
N
N
N
G
G
w
N
o
F
O
\V
N
,—
O
N
\v
w
w
—
O
w
W
O
N
N
o
o
o
N
o
o
N
D
G
D
D
o
N
O
w
w
N
W
W
W
N
w
w
W
w
N
N
N
N
N
W
F
w
N
\v
w
W
W
\V
W
N
\v
w
\\"
G
N
N
N
G
N
N
N
N
G
G
N
N
G
F
F
D
N
G
D
w
F
N
W
F
O
F
\V
N
\Y
NNWWWWWNNN
ONNONN— — NO
left
old
December, January and March and in all four is asymmetrical. The moult is
not basically centrifugal, as stated by Friedmann (1930), nor does it adhere to
the strict 5-1-4-2-3 order described for B. abyssinicus by Stresemann &
Stresemann (1966). It has been found that the latter, the basic bucerotid mode
of rectrix moult, does not hold for a large sample of three species oiTockus
hornbills (Kemp in prep.) but rather the modified system 5 or 1 - 4 or 2 - 3.
This modified mode of rectrix moult may also be more applicable to the data
shown for B. leadbeateri m Table 1.
Active moult is found in specimens taken between November and June
and it is possible that it does not occur from July to October (the main egg
90
laying period) even in adult males and immatures of both sexes. Kemp (in
prep.) reports nest helpers which are doubtless the immature members of the
parties in which this species is habitually seen.
The palate of the subadult female in the National Museum was purplish
brown, the iris was pale yellow and the throat vermillion fringed with yellow.
Other females in the National Museum and Transvaal Museum had violet
blue as well as vermillion on the throat. Two specimens in the Transvaal
Museum (TMP b and e in Table i) appear to have been mis-sexed as males,
whereas the violet blue instead of red on the throat and small si2e (see below)
indicate that they are females. From field observations (Kemp in prep.) it
appears that breeding females, and probably all adult females, have a violet
blue patch on the throat. The age at which the violet blue throat is attained
is uncertain.
Only one violet blue throated (fully adult female), one pink throated
(immature), and one yellow throated (first year) bird, is found in B. leadbeateri
parties which normally number four or five birds (Kemp in prep.) It appears
that these parties may consist of an adult male, an adult female and their
sexually immature offspring which are produced at a maximum rate of one a
year. The normal clutch is two eggs but there is no record of more than one
well grown nestling in a nest. A 50 per cent, mortality among immature
jB. leadbeateri is by no means unlikely. This would mean that a party of five
birds (two adults, three immatures) represents on average the result of six
years of reproduction. Thus it is likely that B. leadbeateri is not sexually
mature until at least its eighth year.
The age at which sexual maturity is attained can also be approached from a
study of the moult of the primaries. Of the 16 specimens we have examined
only one has yet to moult the first of its primaries. Of the remaining 1 5 only
three show active moult. Thus the period in which the primaries are replaced
and which in most birds defines the total moult cycle, must be a very long
one: we suggest at least three years. On this basis simple descending moult
occupies the second, third and fourth years and normal wing length is
attained when the second generation primaries 6 and 7 (the longest ones)
have grown. The first diagonal stepwise moult occupies the fifth, sixth and
seventh years and only after this can the bird be regarded as fully adult in
TABLE 2
Mensural data suggestive of sexual dimorphism in Bucorims leadbeateri
Wing in mm
Source
6*
?
Sexed 6*> t> ut with
violet blue on throat
NMB
550 (juv), 610
450, 510, 550, 560
QVM
580
—
TMP
525 (juv), 536, 558
544
512, 533, 546
DM
—
575
Verheyen (1953)
520 (juv), 545,
570, 570
530
Sanft (i960)
496-618, av. (33)
560
495-5 50, av (10). 538
Culmen in mm
NMB
198 (juv), 210
178, 190, 195, 195
QVM
218
TMP
187 (JUV), 221
207
182, 185, 200
DM
180
Verheyen (1953)
160 (juv), 197,
207, 210
185
Sanft (i960)
190-221, av.
168-215, av.
(30) 207
(10) 192
plumage. During this second moult only females can be separated from
adults through their lack of a violet blue throat. Second moult males are
phenotypically adult and are treated as such in Table i. In nearly all hornbills
immature birds resemble adult males, not adult females. A juvenile in Tables
i and 2 is a bird which has not yet grown its second generation primaries
6 and 7.
It appears from Table 2 that females are mensurally somewhat smaller
than males, especially in the length of the culmen. Wings were measured
with a tape from the carpal joint to the tip of the longest primary which is
sometimes the sixth but more often the seventh. Culmens were measured
from the base to the tip with a tape, apparently also the method used by
Verheyen (1953). Sanft's (i960) ranges do not distinguish between adult and
juvenile birds. This is a pity since it appears from Table 2 that juvenile birds
are mensurally smaller than adults and subadults of the same sex; nonetheless
further study is needed.
The fresh subadult female brought into the National Museum and very
hungry weighed 2,230 g which is over 45 per cent, lower than the female
weight of 4,000 g recorded in Sanft (i960). Weights of males recorded in
Sanft(i96o) and Verheyen (1953) are 3,500, 3,575 and 3,937 g.
We are obliged for facilities for study to M. P. Stuart Irwin in respect of the
National Museum in Bulawayo, to M. A. Raath in respect of the Queen
Victoria Museum and to P. A. Clancey in respect of the Durban Museum.
References :
Friedmann, H. 1930. The caudal moult of certain Coraciiform, Coliiform and Piciform
birds. Proc. U.S. Nat. Mus. 77(7): 1-6.
George, W. G. & Casler, C. L. 1972. Subalular apterium in birds. Auk 89 : 245-262.
Sanft, K. i960. Bucerotidae. Tierreich 76.
Stresemann, E. & Stresemann, V. 1966. Die mauser der Voegel. Journ. Ornith., sonderheft.
Verheyen, R. 1953. Exploration du Pare National de VUpemba. Fasc. 19. Oiseaux. Brussels:
Inst. Pares Nat. Congo Beige.
The Status and Characters of the Races of the
Red-backed Shrike Wintering in the South African
Sub-Region
by P. A. Clancey
Received ptb January 197$
The Redbacked Shrike Lanius collurio Linnaeus is one of the commoner and
more widely distributed of the Palaearctic migrants present in Africa south
of the Cunene-Okavango-Zambesi Rivers during the period of the northern
winter. On its wintering grounds, which extend from Angola, southern
Zaire and Tanzania to the Orange R. and the eastern Cape (Moreau [1972]
believes the winter range in eastern Africa to extend as far north as Kenya
and the southern Sudan, and follows Fry in also listing Nigeria in West
Africa), it occurs in various savanna woodland facies, and is perhaps most
numerous in acaciaveld, in degraded areas, and on the scrubby and over-
grown fringes of primitive cultivation. Redbacked Shrikes commence to
arrive in the Sub-Region in the latter part of October, but do not become
widespread until the second half of November, the majority departing the
first half of April, but with a few laggards still present mid-May, and even
early June in Zambia according to Benson et al. (1971). During their sojourn
92
the birds undergo a complete moult, which is at its height January-mid-
March, the populations commencing their northward movement through
the eastern half of Africa shortly after its completion.
Variation used by workers to arrange the populations into acceptable
subspecies is restricted to males, although, in fact, both sexes show reasonably
well-marked geographical variation. In males, the forehead varies from pale
neutral grey to almost white in far eastern populations, there is appreciable
variation in the shade of grey of the rest of the head-top, hind and sides of
the neck and the upper mantle, while the mantle and scapulars vary from
almost tawny to deep vinous chestnut. In females the upper-parts vary from
dull olivaceous to dark vinaceous brown, this variation appearing to follow
a closely similar distributional pattern to that of the males. On the ventral
surface there is marked variation in the whiteness or buffiness of the ground
and in the degree of pectoral and lateral squamation. Currently three of the
four races recognised by Vaurie (1959) are admitted to the South African
list (Clancey [1966]; S.A.O.S. List Committee [1969]; McLachlan & Liver-
sidge [1970]), nominate L. collurio and L. c. kobylini (Buturlin) supposedly
common and widespread seasonally, with the central Palaearctic L. c.
pallidifrons Johansen known from only two or three records from the north-
ern Cape and South- West Africa. In an effort to arrive at a more satisfactory
assessment of the relative abundance of these races on the South African
wintering grounds and the characters by which they may be determined in
this region, some 200 <$<$ from southern and eastern Africa were assembled
and critically studied. In order to minimize the influence of habitat and
seasonally induced as opposed to genetically based differences on the con-
clusions drawn, the precautionary measure of dry cleaning about a third of
the assembled specimens was taken.
A series of some two hundred males is scarcely adequate for this type of
research, and only from the territories of the Transvaal, Rhodesia and Zambia
were samples anything like satisfactory. In allocating African hibernal
material to races 10 <$<$ L. c. collurio from the western Palaearctic (Sweden,
Lithuania, Germany, Czechoslovakia, Austria and Hungary) were used as
controls. Of L. c. kobylini and L. c. pallidifrons I used short series kindly
determined as of these races by Professor Hans Johansen, formerly of the
Zoologisk Museum, Copenhagen, and the describer of the latter subspecies.
No untoward difficulty was experienced in allocating the great bulk of the
material to definite subspecific categories. Conclusions finally reached were
on the basis of series laid out and viewed facing the light source, the sky
moderately overcast and the time mid-morning.
Bearing in mind the barely adequate nature of the material presently
available, the following tentative conclusions were reached: (i) L. c. pallidi-
frons is as numerous or slightly more so than L. c. collurio. Of 146 £<§ from
the South African Sub-Region studied 3 9 per cent, were pallidifrons, with
some 35 per cent, nominate collurio. (ii) Contrary to expectations, L. c.
kobylini is far less numerous than either L. c. collurio or L. c. pallidifrons (26
per cent.); (iii) L. c. pallidifrons winters largely in the Kalahari and adjacent
xeric areas of the interior of the Sub-Region; (iv) L. c. kobylini winters
extensively in northern South- West Africa, and in the east from Rhodesia
and Mozambique south to Natal; (v) on the basis of present data no hibernal
centrum is definable for L. c. collurio, which occurs throughout. In so far as
the small British Isles population (L. c.juxtus Clancey), with backs in males
umber brown rather than rufous, is concerned, this study shows that it
93
does not extend to within present limits, and its wintering grounds are
clearly extra-limital (?=hygric regions of Angola or southern Zaire).
In the case of Zambia, the one extra-limital African territory within the
species' wintering range from which a reasonably adequate panel of adult
males has been available to me, all three races wintering in the South African
Territory
Cape
N
IO
TABLfc
L. c. collurio
5 (50%)
3 Nov. - 15 Jan.
1
Race
L. c. pallidifrons
4(40%)
Jan. - 27 Mar.
L. c. kobylini
1 (10%)
12 Mar.
South-West
Africa
9
2(22-2%)
8 Dec. - 6 Mar.
i(ii-i%)
3 Dec.
6(66-7%)
9-28 Mar.
Botswana
17
6(35-3%)
13 Nov. - 30 Mar.
9(52-9%)
12 Nov. - 15 Apr.
2(11-8%)
20 Feb.
Rhodesia
52
15(28-8%)
14 Nov. - 17 Mar.
20(38-5%)
23 Nov. - 10 Apr.
17(32-7%)
1 1 Nov. - 1 1 Apr.
Transvaal
(mainly west)
35
14(20%)
7 Nov. - 12 Apr.
17(48-6%)
12 Nov. - Apr.
4(ii-4%)
28 Sep.(?) - 23 Feb
Natal, Swaziland
& East Griqualand
18
6(33-3%)
19 Nov. - 11 Feb.
6(33-3%)
24 Nov. - 4 Mar.
6(33-3%)
4 Jan. - 1 Apr.
Mozambique
5
3
27 Nov. - 24 May
—
2
1 Mar. - 7 Apr.
Extra-limital
Zambia
35
ii(3i-4%)
27 Oct. - 7 Apr.
16 (45-7%)
29 Oct. - 14 Apr.
8(22-9%)
21 Mar. - 8 Apr.
Lanius collurio Linnaeus
Allocation of South African Sub-Region and Zambian material of adult £$ Redbacked
Shrikes to subspecies. The assumed relative abundance of each subspecies in a territory's
sample is given in the form of a simple percentage. The first and last dates of specimens in
each sample are enumerated for the guidance of future workers.
Sub-Region were found to occur, as was to be expected. It will be appreciated
from accompanying Table I that the same position as recorded for South
Africa obtains in Zambia, with pallidifrons the commonest of the three
Redbacked Shrike races, and kobylini the least so. It is evident from the com-
ments of Benson et al. (1971), that Zambian data in large measure corrobor-
ate my findings for the South African Sub-Region.
Vaurie (1959) has defined the breeding ranges and racial characters of the
four races of L. collurio, sens, strict. , on the basis of material from the Palae-
arctic. Examination of skins of both sexes just through the moult from South
Africa shows that the criteria ascribed to the individual races by Vaurie
require to be modified and expanded somewhat. Hereunder I give my con-
clusions as to the characters of use in allocating wintering material taken in
southern and eastern Africa to subspecies. The breeding ranges have been
laid down in Vaurie's work; in so far as Africa is concerned, the races
appear to be synhiemal, or mainly so.
(a) Lanius collurio collurio Linnaeus, 1758: Sweden
o*. Forehead Pale Neutral Gray (Ridgway [191 2], pi. liii), merging
insensibly in to Neutral Gray (same pi.) on the crown, nape, hind and
sides of the neck and upper mantle; mantle and scapulars Snuff Brown
94
(pi. xxix). Tertials with snuff brown extending over most of outer and
part of inner vane.
$. Upper-parts Saccardo's Umber (pi. xxix), the hind neck greyer, and
breast and lateral ventral surfaces heavily squamated with dusky on a
whitish ground.
Remarks: Specimens unequivocally of this subspecies were examined from
South- West Africa and the Cape, east to the Mozambique littoral, with no
evidence from the available data of a preference for any particular sector of
the Sub-Region.
(b) Lanius collurio pallidifrons Johansen, 1952 (1944): Tomsk, Western
Siberia, U.S.S.R.
o*. As in nominate collurio, but with the grey of the forehead lighter,
often whitish (Pallid Neutral Gray [pi. liii]), merging to Light Neutral
Gray (same pi.) on the crown, nape, neck and upper mantle, this sharply
demarcated from the redder mantle and scapulars (close to Russet [pi.
xv]). Tertials as in collurio.
9- Upper-parts rather lighter, slightly redder, less olivaceous, especially
over the mantle and scapulars than in collurio, and hind neck greyer
(mantle and scapulars Sayal Brown [pi. xxix]). Ventrally about the same.
Remarks: Originally identified from South Africa on the basis of a male
taken at Riverton, Kimberley, northern Cape (Clancey [1961]), and later
recorded from Olyvenhout Drift, near Upington, on the Orange R. by
Winterbottom (1968) and "Quickborn" Farm, Okahandja, South- West
Africa (identified specimen in Transvaal Museum not alluded to by Winter-
bottom [1971]). Now established as almost certainly the most numerous of
the three subspecies occurring within present limits, wintering largely in the
Kalahari and peripheral xeric areas of the South West Arid District. Records
are now available from the northern Cape, South- West Africa (Okahandja),
Botswana, Rhodesia, the Transvaal (mainly dry western) and Natal (coast).
There are also numerous records from Zambia, in which territory probably
largely a transient to and from desertic areas further south.
A typical example of pallidifrons from Kitale, Kenya, is in the collection
of the South African Museum, Cape Town. The specimen concerned is
dated 7th April, 1929, and was presumably on northbound migration when
taken.
Vaurie (1959) records that both pallidifrons and nominate collurio hybridize
freely with/., c.phoenicuroides (Schalow), 1875 : Chimkent, Russian Turkestan,
where their ranges meet, though there is no evidence of any stage of this in
South African material (of adult males, as well as sub-adults and adult
females), and such interracial hybrids must winter within the Ethiopian non-
breeding range oiphoenicuroides, which is from L. Chad, east to south-western
Arabia and Somalia, south in the east of Africa to Malawi (Livings tonia).
Not south of the Zambesi R.
(c) Lanius collurio kobylini (Buturlin), 1906: Kutais and Ssuran, Trans-
caucasia, U.S.S.R.
<£. Grey of head-top, hind and sides of the neck and upper mantle darker
and bluer than in nominate collurio (about Dark Gull Gray [pi. liii]),
this frequently more extended caudad over the mantle; mantle and
scapulars darker and more saturated, being deep vinous brown (Chest-
nut Brown [pi. xiv]). Tertials darker, with the snuff brown generally
restricted to the lateral outer vane, and the tip starkly white or huffish
white in each feather when in a pristine condition.
95
(
2 6 j UN19t*
$. When freshly moulted much darker vinaceous brown above than
collurio (mantle and scapulars Prout's Brown [pi. xv]). Below with
buffish or vinaceous tinge to ground,* and reduced, often vestigial,
squamation.
Remarks: This race appears to winter extensively in northern South- West
Africa, and in the east in Rhodesia and Mozambique, south to Natal; it is
relatively poorly represented in the samples from the northern Cape (i),
Botswana and the western Transvaal, in which xeric region pallidifrons pre-
dominates. L. c. kobylini occurs commonly on. passage in coastal Kenya, being
the only race represented in the series from that area in the Durban Museum
collection. Zambian records are dated 21st March-8th April, suggesting its
occurrence there is also that of a transient.
AC KNOWLEDGEMENTS
For the loan of material to augment that in the Durban Museum, I
acknowledge indebtedness to the South African Museum, Cape Town (Dr.
J. M. Winterbottom), the Transvaal Museum, Pretoria (Mr. A. C. Kemp),
the State Museum, Windhoek (Mr. P. J. Buys), and the National Museums
of Rhodesia, Bulawayo (Mr. M. P. Stuart Irwin). I am also grateful to Pro-
fessor Hans Johansen, formerly of the staff of the Zoologisk Museum,
Copenhagen, for his assistance with these shrikes and for determining
material submitted to him. Some two hundred adult males and a rather
larger number of sub-adult males and females were studied during the course
of this investigation.
References :
Benson, C. W. et al. 1971. Birds of Zambia. London: 293.
Clancey, P. A. 1961. The West Siberian race of the Red-backed Shrike in South Africa,
Ostrich, 32 (3): 143.
— 1966. A Catalogue of Birds of the South African Sub-Region, Durban Mus. Novit., 7
(12): 530, 531.
McLachlan, G. R. & Liversidge, R. 1970. Roberts' Birds of South Africa. Cape Town: 481.
Moreau, R. E. 1972. The Palaearctic- African Bird Migration Systems. London and New York:
130-132.
Ridgway, R. 191 2. Color Standards and Color Nomenclature. Washington.
S.A.O.S. List Committee, 1969. Check List of the Birds of South Africa. Cape Town: 272, 273.
Vaurie, C. 1959. Birds of the Palearctic Fauna. London. 1 (Passeriformes) : 96-100.
Winterbottom, J. M. 1968. Check List of the land and fresh water birds of the western Cape
Province, Ann. S. Afr. Mus., 53 (1): 232.
— 1 97 1. Preliminary Check List of the Birds of South West Africa. Windhoek: 215, 216.
ADDENDUM
Since the above report on Lanius collurio in the South African Sub-Region
was prepared and submitted for publication, I have been able to study a
recently obtained series of freshly moulted examples from Angola. Six $$,
two $$ in immaculate dress from Mocamedes (Bibala) and Huila (Chibemba
[Gambos] and Cahama) taken between 19th February and 6th April (1967-
1970) are all unequivocal L. c. kobylini. These specimens have wings in $$
94. 5 - 9 8 > $$ 94- 5, 95 • 5 mm; weights <$$ 27 - 32, $$ 24, 25 gm. The dis-
covery that L. c. kobylini winters in numbers in south-western Angola ties
in with the finding enunciated above that it is the main race occurring in
northern South- West Africa, immediately south of Mocamedes and Huila.
I am grateful to Dr. A. A. da Rosa Pinto for the loan of the material in the
collection of the Instituto de Investigacao cientifica de Angola, Sa da
Bandeira.
96
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Published by the BRITISH ORNITHOLOGISTS' CLUB and printed by
The Caxton and Holmesdale Press, 104 London Road, Sevenoaks, Kent.
Bulletin of the
British Ornithologists' Club
Edited by
C. W. BENSON
Volume 93 No. 3
September 1 973
Committee
Sir Hugh Elliott, Bt., o.b.e. {Chairman)
J. H. Elgood {Vice-Chairman)
C. W. Benson, o.b.e. {Editor)
R. E. F. Peal (Hon, Secretary)
P. Tate {Hon. Treasurer)
J. H. R. Boswall
Dr. P. J. K. Burton
P. Hogg
C. J. Mead
Bulletin of the
BRITISH ORNITHOLOGISTS' CLUB
Vol. 93 No. 3 Published: 20 September, 1973
The six hundred and eighty-third meeting of the Club was held at the
Cafe Royal, 68 Regent Street, London, W.i., on Tuesday, 17th July, 1973
at 7 p.m.
Chairman: Sir Hugh Elliott, Bt., o.b.e., present 12 members and four
guests.
Captain Collingwood Ingram read the communication given below and
exhibited down feathers of an Eider So materia mo//issima*.
Mr. Robert Spencer gave an address on feeding by Siskins in gardens,
illustrated by slides.
*"Fifty-three years ago, at one of the Club's dinners {Bull. Brit. Orn. CI.
40, 1920: 121), I showed an exhibit which aroused so much interest that I
thought some of you might like to see it. While on a visit to one of the more
northern of the Orkney Islands I was able to examine a fully developed
embryo of an Eider. It was in an egg which had been abandoned by the
incubating bird when on the point of being hatched. The dead embryo was
still wet from the viscous fluid of the egg's albumen, and so had the usual
bedraggled appearance of an unhatched chick. What at first sight seemed to
be a clotted mass of wet hairs proved on close examination to be in fact the
chick's covering of down feathers, unrecognisable because every feather was
enclosed in a protective sheath. Although the encased neossoptiles, or down
feathers, each had about twenty barbs, these were so tightly compressed
within the protective sheath that they looked more like single horse-hairs
than feathers. When dry the integument of the protective sheaths becomes
very friable, and in an incredibly short time after hatching it disintegrates
from the bottom upwards, finally disappearing completely. Thus in a matter
of minutes the erstwhile besodden and bedraggled object becomes trans-
formed into a fluffy little duckling clothed all over in a dark blackish down".
Migration of Motacilla alba alha
by John Larmuth
Received rph April, 197)
During the last week of March 1962 I was on board the Adriatica Lines M.V.
Esperia bound for Beirut, from Naples, via Alexandria. The second part of
this voyage is on an approximately north-easterly course. At 19.30 hrs.,
when night had fallen and the ship was between 100-150 miles north of the
Egyptian coastline, it was noticed, upon coming on deck, that the air within
visible range of the ship's lights was occupied by a dense flock of wagtails of
the genus Motacilla flying parallel to the ship at a speed 2-3 knots greater than
that of the ship (about 17 knots). It was estimated that a minimum of 5-
10,000 birds were passing the ship at any one time and that any one bird
took about two minutes to pass from stern to bow. Birds were not observed
returning to the neighbourhood of the ship once they had passed ahead into
the darkness. Every available bit of deck space, and every perch was occupied
by resting birds. The passage was observed until 02.00 hrs., when I retired
to my cabin.
97
In the morning no living birds were seen, but many thousands of dead
Motacilla a. alba were being hosed into the scuppers during cleaning up
operations.
The total number of birds passing the ship during the time of this observa-
tion may conservatively be estimated at 1,000,000.
During flight, in the normal undulatory manner, it was noticed that large
numbers of the birds in the lower part of the flock were flying into waves,
when a water wave crest coincided with a flight trough. It is possible that
the flock was flying at an unusually low height due to the presence of the ship.
[K. D. Smith, to whom the above note was shown, has commented as follows: "I cannot
recall a published record of visual M. alba passage across the Mediterranean on such a vast
scale, though there may be comparable observations in the literature. But it seems reason-
able to suppose that after the desert crossing the migrants ran into strong, southerly
khamsin winds, '. . . which constitute a special hazard to migrants. They blow with
devastating violence, typically for periods of two or three days, and at their worst cause
sand-storms.' (Moreau, 1961. Ibis 103a: 386). Maybe exhaustion caused them to fly into the
water, and it is difficult to account for the mortality other than turbulent conditions further
south which may have prevented a landing on the coast of Egypt." — Ed.\
Wing moult of Ruddy Ducks in Manitoba
by W. Roy Siegfried
Received jth May, 19J3
Two southern hemisphere stifftails, Oxyura australis and O. maccoa have been
reported as moulting the remiges twice annually (Frith 1967; Siegfried 1970).
This paper presents evidence for a similar double wing moult in a wild
population of the northern hemisphere Ruddy Duck O.j.jamaicensis.
During May - August 1971 I collected Ruddy Ducks on their breeding
grounds in south-western Manitoba, Canada. Seventy specimens were
examined for moult. I also noted the presence or absence of wing moult in a
large number of live Ruddy Ducks observed in the field. Additional, useful
information was obtained from birds which had been live- trapped.
TABLE 1
Numbers of male and female Ruddy Ducks replacing remiges. Totals of birds checked for
presence or absence of wing moult are given in parentheses.
May June July August
1-15 16-31 1— 1 5 16-30 1— 1 5 16-31 1— 1 5 16-31
St
(38)
(59)
(65)
(42)
(79)
4
(47)
21
(60)
9
(22)
99 1
(43)
(61)
($0
(45)
(62)
(60)
(6 2 i)
2
(9)
Two out of 22 specimens examined in the hand during the period 15th
May to 1 5 th June were found to have the bases of the fully grown primary
shafts still encircled by remnants of the feather sheaths. In two of these birds,
male and female both in full adult breeding plumage, the remnant sheaths
extended 5-10 mm up the shafts of the distal primaries. The primary feathers
of all specimens collected in May were fresh and immaculate. They appeared
to have been newly acquired, probably in late-winter/early spring, and prior
to the birds' arrival in early May in Manitoba. Two females, however, were
98
found to have developing, less than half-grown remiges. Clearly, these
individuals must have commenced moulting the wings after having com-
pleted the spring migration. One of these birds was first observed on 15 th
May, and the other individual was taken in a decoy trap on 9th June. This
female was fitted with a beak marker. Subsequently, she made one attempt
to breed, but lost the eggs to a predator. I consider these two cases, of birds
moulting soon after arrival on the Manitoba breeding grounds, as atypical ;
and believe that the birds were first-year individuals which hatched relatively
late in 1970 and/ or in an area to the south of Manitoba. Table 1 shows that,
commencing August the incidence of moult of the remiges was common,
especially in males, in the population. Those males which were in the process
of replacing their remiges were still in mainly bright breeding plumage.
In summary, it appears that generally Ruddy Ducks breeding in Manitoba
moult their wings in late-summer/early-fall before departing for their winter
quarters where presumably they undergo a second wing moult in late- winter/
early-spring before migrating northwards.
In a previous paper (Siegfried 1970), I could offer no explanation as to
what the selective advantage of this remarkable double wing moult might be
to the Oxyurini. I can add very little to this except to note that none of the
Ruddy Duck specimens, including those collected in late summer, showed
any obvious signs of hard wear on the remiges. By contrast, a male O.
maccoa examined during its breeding cycle in the south-western Cape, South
Africa, had its secondary flight feathers (nos. 1 - 6) worn down to about
half their normal length. Wear of this kind presumably might result by
frequent performance of display flights across water (see Johnsgard 1965,
1968 for descriptions of this behaviour). However, display flights occur less
often in O. maccoa than in O. j. jamaicensis, though the season for their
occurrence is considerably longer in individual O. maccoa (Siegfried unpubl.)
Acknowledgements :
The work was made possible by grants from the Delta Waterfowl Research
Station, the Chapman Memorial Fund of the American Museum of Natural
History, the South African Council for Scientific and Industrial Research
and the University of Cape Town. I am grateful to these bodies for their
support. The Canadian Wildlife Service gave permission for the taking of
birds.
References :
Frith, H. J. 1967. Waterfowl in Australia. Sydney: Angus & Robertson.
Johnsgard, P. A. 1965. Handbook of Waterfowl Behaviour. London: Constable.
— 1968. Some observations on Maccoa Duck behaviour. Ostrich 39: 219-222.
Siegfried, W. R. 1970. Double wing-moult in the Maccoa Duck. Wildfowl 21: 122.
Numenius rninutus, Falco subbuteo and
Caprimulgus europaeus in the Seychelles
by C. J. Feare
Received 16th May, 19 jf
Among many migrant land and shorebirds recorded in the Seychelles in
October, November and December 1972 were some new to the Malagasy
Region. Three of these, Numenius rninutus, Falco subbuteo and Caprimulgus
europaeus, were identified from photographs by Messrs. C. W. Benson and
99
R. Wagstaffe. The photograph of N. minutus was taken by Dr. R. J. Raines,
while those of F. subbuteo and C. europaeus were taken by Mr. Guy Savy, and
all of these have been deposited in the University Museum of Zoology,
Cambridge. These species all occurred on Bird Island, the northernmost
island of the Seychelles archipelago. The N. minutus was first seen there on
14th October 1972, during my residence on the island, and I therefore had
many opportunities of watching and filming the bird. According to Mr. Savy,
it was still present in late December 1972. The F. subbuteo was found dying
and the C. europaeus found dead in early December by Mr. Savy, who
measured the wing (flattened chord), bill (from feathers) and tarsus of each
bird, and kindly sent photographs of them to me.
The following details of these birds were obtained :
Numenius minutus:
Inflight, this bird was immediately distinguishable from all other palaearctic
shorebirds by its resemblance to a small, sandy-coloured Whimbrel iV.
phaeopus, but completely lacking white on the rump and back. Its general
appearance was of a warm sandy-buff coloured bird with darker brown
markings, especially on the dorsal surfaces. It lacked the "deep-chested"
appearance of a Whimbrel.
At rest, it was much smaller than a Whimbrel, and also smaller than a Bar-
tailed Godwit Limosa lapponica. It stood about as tall as a Grey Plover
Squatarola squatarola but had a much slimmer build. The back and wings were
creamy-buff spotted with dark brown. The tail was similarly marked, but the
spots tended to form bars across the tail. The rump was the same colour as
the back. The breast was sandy-buff, almost clear but lightly streaked with
brown, especially on the sides. The belly and under tail coverts were a
clear creamy-buff, and long creamy-buff feathers projected backwards from
the top of the thigh. The superciliary stripes, a stripe down the centre of the
crown, and the cheeks, were creamy-buff, much paler than in Whimbrels,
while the rest of the crown and a mark between the superciliary stripe and
the cheek were rich brown. The back of the head had a much more square
appearance than in Whimbrels. The bill was comparatively short but down-
curved ; it was brown with a pink base, this being especially noticeable on the
lower mandible. The legs were creamish-grey.
When flushed the bird uttered a fairly faint but harsh croak, sometimes
di- or tri-syllabic, in which case it resembled the beginning of a Whimbrel's
call. The only other call heard was a trill, which was given while Dr. Raines
was stalking the bird in order to obtain photographs.
It was always found in open country on the island, usually on the half-
completed airstrip, but occasionally in grassy areas in the Sooty Tern Sterna
fuscata colony.
This species breeds in north-east Siberia and normally winters in Austra-
lasia, and its appearance in Seychelles was probably a freak occurrence.
Falco subbuteo :
This bird was captured and photographed alive in early December 1972,
but it subsequently died. The measurements obtained were: wing 265 mm,
bill 14 mm, tarsus 26 mm. According to Mr. Benson the bird was an adult,
and the wing length places it in the nominate race.
Between October and April of the years 1971-72 and 1972-73 several
unidentified falcons have been seen in the Seychelles. Benson & Penny {Phil.
Trans. R. Soc. 260B, 1971: 515-516) have stressed the difficulties of identi-
fying falcons on Aldabra, where they considered that most migrants were
probably attributable to F. eleonorae or F. concolor. This may also be true of
migrant falcons in the Seychelles, but F. subbuteo may also occur regularly.
Caprimulgus europaeus:
This bird was found dead in early December 1972, and its measurements
were as follows: wing 190 mm, bill 6 mm, tarsus 15 mm. In view of its dark
colour and long wing Mr. Benson concluded that it was of the nominate race,
and the absence of white on the outer rectrices and of white spots on the
primaries indicated that this bird was a female.
Subalpine and Grasshopper Warblers in Sierra Leone
by G. D. Field
Received 10 th May, 1973
Neither the Subalpine Warbler Sylvia cantillans nor the Grasshopper Warbler
Locustella naevia has hitherto been recorded wintering in the forest zone of
West Africa. Moreau (1972) summarised the available information on the
wintering grounds of S. cantillans as the borders of the southern Sahara
eastwards to Lake Chad, with the most southerly record at about n°N in
Nigeria, while for L. naevia, apart from one record of L. n. straminea in
Ethiopia, there are now several records of spring and autumn passage from
the western Sahara between Morocco and Senegal, from which Moreau
concluded "It is at least certain that many Grasshopper Warblers must winter
in West Africa, presumably in a zone more humid than Senegal". It may,
therefore, be of interest to record the occurrence of both species in Sierra
Leone in the early months of 1973.
On the evening of 1st February at 18.45 (almost dusk) I saw a small bird
fly across a creek of the Sierra Leone River about eight miles south-west of
Freetown at 13 08' W and 8° 21' N. It disappeared into an isolated triangular
patch of Avicennia mangroves, only a few yards thick, growing on a piece of
waste land beside the jetty. Idly wondering whether it might be an Acro-
cephalus warbler, I approached and caught a glimpse of a mantle so blue-grey
that I was reminded of one of the grey Muscicapa flycatchers, but behaviour
showed at once that it was a warbler. By walking slowly back and forth
along the two landward sides of the mangrove thicket I was able to catch
sufficient fleeting views to take an adequate description. The bird was not
shy but extremely active, insect-hunting low down among the leaves, rarely
more than two feet above the ground though often close to the edge. Often
its progress could only be followed by the movements of the leaves as it
worked through the bushes ; once it made a little vertical flight up and down,
flycatcher-like. I watched it until it became too dark for further observations
and took the following description : upper parts blue-grey, particularly bright
on the head, wings browner with pale feather edgings ; a clear and striking
white moustache streak; underside brick red or pinkish russet, becoming
duller on belly, under tail coverts whitish. Beak dark, eye red (this would, in
fact, have been the eye-ring).
The isolated occurrence of one Subalpine Warbler is not of great signifi-
cance, though it is interesting that the habitat — Avicennia mangroves — is the
same as that recorded in the Gambia. By its very nature such a habitat is not
easily searched for birds and there are vast areas between Gambia and Sierra
Leone which could be frequented by the species without discovery.
The record of the Grasshopper Warbler is more interesting in that at
least two and possibly three individuals were concerned. I spent March 25 th
to 29th at a small town, Njala Komboya, in central Sierra Leone, ii° 27' W
and 8° 12' N, for the purpose of making a rapid bird survey of an area not
previously visited. The country is remote and entirely rural. A barely
motorable road reaches Njala from the south and beyond are only footpaths
to villages from which agriculture is practised on the basis of shifting culti-
vation. It is an area of well- watered, often swampy, valleys and small hills
rising at their highest to 1,400 ft. a.s.l. All was once forested and patches of
high forest remain, particularly around villages as 'sacred bush', as well as
riverine forest along streams. The bush is cut and burnt for a season's
agricultural operations and then left to regenerate for several years, the
result being a thick and virtually impenetrable tangle. Other areas have been
taken over by grass and here a variety of savanna species have entered, there
resulting a typical mosaic of forest and derived savanna vegetation.
I was at once struck by the richness of the bird fauna, particularly of
passerine palaearctic migrants: Sylvia borin was extraordinarily abundant
while Hippolais polyglotta and Acrocephalus scirpaceus were seen far more often
than normally in Sierra Leone and I suspect that the birds were on passage
especially as the commonest migrant warbler Phylloscopus trochilus was vir-
tually absent and had presumably already left. The Reed Warblers were
particularly interesting, seen not only in their normal wintering habitat of
long, coarse grass but also in burnt vegetation and even thick bush.
One north-facing hillside consisted at the top of thick, dry bush some 25
feet high, on one side of the path newly felled. Down the slope there was a
patch of rocky deciduous forest, dominated by a few huge Ceiba pentandra
trees, the smaller trees beneath almost bare and with little ground vegetation.
Below this was lusher thick bush up to 1 5 feet high and along the valley floor
ran a stream with a strip of dense riverine forest along its banks, the whole
slope being some five or six hundred yards long. On 28th March in mid
morning a small warbler flicked across the path at the lower edge of the forest
patch and crept at great speed through the tangles. Though puzzled I could
only assume that it was a Reed Warbler and as there was no hope of seeing
it again I ignored it. On 29th March at 8 a.m. in the thick bush just above the
stream, some two hundred yards further down, I flushed a small bird from
on or near the ground close beside the path; it flew a few feet and then
skulked through the shrubs close to ground level and crept away. All I caught
was a glimpse of a brown upperside heavily spotted and noted "very grass-
hopper warbler-like". I waited for some time without seeing it again but a
few minutes after it had disappeared I heard a short, sharp, truncated burst
of reeling song, lasting only a few seconds, quite unlike anything previously
heard in Sierra Leone but reminding me at once of a Grasshopper Warbler.
Two hours later on my way home I flushed another bird in the dry bush
at the top of the hill, again from close beside the path. Because it was between
the path and the felled area the bird could only fly three or four yards from !
me, where it stopped on a small branch two feet up and peered at me, dipping
forward and flicking from side to side in undecided fashion, later taking little |
hops behind the leaves and stopping again to peer, sometimes craning right I
up to look at me over the leaves, appearing in such a posture almost pipit-like, I
Finally, having made up its mind, it threaded its way quickly and efficiently
through the vegetation, coming in fact extremely close to me but most of j
the time lost to sight behind the thick foliage of a felled tree, and when I
moved to keep pace with it it disappeared entirely.
Despite the fact that it was moving almost the whole time I had good views
at close range and took the following description: general appearance
distinctly smaller than Sylvia borin, slimly proportioned with a rather long
and wedge-pointed tail, beak straight, neither pronounced like Acrocephalus
scirpaceus nor curved like the Cisticola genus. When moving it crept hunched
and mouselike, though when craning to look at me it stood quite tall. Whole
bird rather pale olive-brown ; head top finely streaked with no noticeable eye-
stripe but a definite small circle of pale bare skin round the eye, mantle
rather plain olive-brown with fewer spots but back and wings heavily
spotted with dark brown, flight feathers dark. Tail plain brown with a slight
suspicion when fluffed out of a pale outer edge. Underside : chin and belly
whitish, breast pale olive-brown, faintly drop streaked, flanks and under tail
coverts far more heavily streaked. Eye dark, beak upper mandible dark,
lower rather bright yellow (it was difficult to be certain how the two colours
combined, the impression being that most of the yellow was at the gape
rather than the tip), legs rather bright pinkish flesh. The whole bird gave the
appearance of being decisively coloured and newly moulted.
There can be little reasonable doubt that two individuals were involved on
29th March and possibly a third on 28th March. That such a skulking bird
should be seen twice in one morning suggests that it was locally abundant
and this, in conjunction with the movements of other warblers in the area,
suggests that some migratory movement was taking place.
Reference :
Moreau, R. E. 1972. The Palaearctic- African Bird Migration Systems. London & New York:
Academic Press.
Some calls and behaviour patterns of the Plumbeous
and Rufous Pigeons
by Derek Goodwin
Received 21st May, 197)
INTRODUCTION
In October and November 1972 I made some observations on Plumbeous
Pigeons Columba plumbea in the Serra dos Orgaos National Park and in the
Boraceia Forest Reserve, in south-eastern Brazil, and on a captive Rufous
Pigeon C. cayennensis in a large aviary at the Belem Zoo. With the exception
of the advertising coo of the Rufous Pigeon, the calls heard and behaviour
patterns observed do not appear to have been described for these species.
They are, therefore, given here and comparisons made with some other
neotropical species (Goodwin 1964, Skutch 1964).
In attempting to transcribe cooing sounds, I have, as previously (Goodwin
1956a, 1970) used 66 to indicate a sound similar to the 00 in the southern
English pronunciation of the words "Rook", "cook" and "look", and 06
for a longer sound similar to the 00 in the words "boot", "root" and "cool".
The sketches are an attempt to give an impression of some of the postures
described. They have no pretentions to art or detailed accuracy.
THE PLUMBEOUS PIGEON
At Serra dos Orgaos this species appeared widespread, although not very
numerous, in tall, dense hill forest and relatively most abundant on areas of
103
high, rocky ground where the largest trees were, with a few exceptions,
smaller in size, and where the trees were interspersed with large succulents
and other shrubs. In this favoured habitat the nearest calling individuals
appeared to be about 400 yards from each other. At Boraceia, where there
was none of this rocky habitat in the hill forest where I observed it, the species
appeared to be much less numerous and far more wary of man. At Serra dos
Orgaos, most individuals permitted approach to within quite reasonable
binocular range without concealment, and with a little stalking and wood-
craft could sometimes be observed from very close quarters. In both places,
however, the denseness of the vegetation often prevented me from seeing
pigeons that were calling very close at hand.
The most detailed observations were made at Serra dos Orgaos on 21st
and 25 th October. Here the behaviour shown suggested that the pigeons
were probably starting their breeding season and that the majority of calling
individuals were males as yet unpaired, as they were usually alone. Further
observations are needed to confirm this, as it is possible that there may have
been undiscovered nests and, if so, that either the female sits for longer
periods than is usual for those species of pigeons in which the share of the
sexes in incubation is known, or that females of this species give the adver-
tising coo in their "off duty" periods.
Plumbeous Pigeons were seen feeding on only two occasions, once a
presumed pair and once a single bird, taking fruit from the canopy of smallish
trees. Each bird satisfied itself in less than 15 minutes, which suggests that,
at this period, these pigeons needed very little time to obtain food.
Voice: The advertising coo is a rather bubbly-sounding "66k, ookook,
ookoo". Unlike most other pigeons whose voice is known to me, C. plumbea
appeared never to repeat the advertising coo two or more times without
pause. One that I repeatedly timed took just under three seconds to utter the
cooing phrase and, when calling regularly, paused for five to six seconds
before calling again. At fairly close range the first note of the phrase was
usually distinct and sounded as loud as the rest but it appeared to carry less
well and at a distance the advertising coo often sounded four-syllabled. One
bird (only) omitted the first note on two occasions. It would, therefore, be
unsafe to assume that the number of syllables in the advertising coo could
alone distinguish this species from the related Ruddy Pigeon C. subvinacea,
whose advertising coo is described as four-syllabled (Skutch 1964).
As with other pigeons, there were individual differences in the advertising
coo that were usually only just noticeable when two birds were repeatedly
calling in sequence. Two individuals, however, habitually uttered more
noticeably divergent advertising coos: one had a hoarse tone in the final
"56" of the phrase, the other considerably lengthened and upwardly stressed
the final note.
Copy of a sound spectograph, made by Dr. D. W. Snow, of the advertising coo of a
Plumbeous Pigeon at Serra dos Orgaos.
104
The advertising coo of the Plumbeous Pigeon appeared to be given in
situations comparable to those of other species (Goodwin 1956a, 1970). It
was heard most often from a single bird perched alone in a tree, sometimes
on a bare or partly bare terminal branch and sometimes within the canopy.
Twice it was heard from individuals whose presumed mates were perched
within a few feet of them and once from a bird sitting side by side with its
mate, in physical contact. It was once heard given as an apparent contact
call between two paired or pairing birds, and I noticed that the presumed
female's calls did not sound less loud or less musical than the presumed
male's.
A single "croorr", with a purring or snoring tone and very variable in
loudness, was frequently heard but usually from birds that were out of sight.
One bird, seen at close quarters, spent about ten minutes preening, then
looked suddenly alert, uttered this call, gave it again about six seconds later
and then started to utter the advertising coo. After a session of cooing it
again preened, then flew to another perch nearby, gave the purring "croorr"
twice, as before, and then again started giving the advertising coo. Later
another individual uttered loud, repeated "croorrs", interspersed with
advertising coos. When I had got within sight of it, I saw the bird was alone
on its perch. It then took wing, performed a display flight, and alighted
elsewhere.
On the morning of 1 ith November, at Boraceia, a Plumbeous Pigeon, "A",
was heard giving the advertising coo from its perch at the top of a dead tree
in dense hill forest; a second bird ,"B", appeared to be answering it with the
same call from a large, bromeliad-covered tree about 400 yards away on the
opposite slope (I was on a path between them). After about a minute, "A"
flew and alighted on a mass of epiphytic growth on a bare tree top about 50
yards from the tree in which "B" was calling. As soon as "A" alighted there,
"B" began to utter the purring "croorr" note almost continuously. At
intervals, for about seven minutes, "A" gave advertising coos and "B"
uttered the purring calls, only breaking off on four occasions to give an
advertising coo. Twice when it did so it omitted the first syllable of the
phrase; there was no doubt about this as on the other two occasions when
the "normal" advertising coo was uttered, the first note was loud and clear.
Then "A" flew to "B", or at least to "B"s close vicinity (I could not
see them), and at once there were emphatic, repeated purring calls but no
sound of fighting. It seemed therefore most likely that "A" had joined a mate
or prospective mate on a potential nest site and that, as well as being used as
an alternative advertising coo, the purring or snoring "croorr" is also the
nest call. In at least two other pigeons, the Diamond Dove Geopelia cuneata,
and the Picazuro Pigeon Columba pica^uro, a call with a snoring or growling
tone is used in both these contexts (Goodwin 1964, 1970).
Some behaviour patterns : The display flight was seen only twice, although I
think that under comparable conditions it would have been seen very many
times from any of the European species of Columba or Streptopelia. I saw it
first from a bird that for the previous 27 minutes had been perched on a bare
terminal bough, uttering the advertising coo at the usual intervals. It sud-
denly sleeked down its plumage, looked alertly about, then towered up at a
steep angle in flapping flight, flattened out and flew at a slight downwards
incline to another perch about 300 yards away, where it soon started calling
again. The second display flight, seen from a bird (mentioned above) that
had been uttering both the advertising coo and the purring call, was similar.
105
In neither case were there audible wing claps, such as occur in high intensity
display flights of all other pigeon species whose display flights are known to
me. Possibly these two display flights of C. plumbea that I saw were at low
intensity but I had the impression that the situation was rather that in this
species the threshold for performing the display flight was relatively high.
It seems possible that this species may, in an evolutionary sense, be in process
of losing this display.
i. Plumbeous Pigeon giving adver-
tising coo.
Plumbeous Pigeon in head back
phase of the flight-intention
movement.
Plumbeous Pigeon parading.
4. Rufous Pigeon nodding.
When making flight-intention movements the Plumbeous Pigeon pulls its
head far back onto its shoulders and then throws it forward with a rather
circular motion. The movement is very similar to that of the Grey-fronted
Dove Leptotila ruf axilla, and the White-bellied Dove L.jarnaicensis, and (if I
recall it rightly) identical to that of the Picazuro Pigeon. When a Plumbeous
Pigeon was alarmed by the sudden swooping of some swifts Strep toprocne sp.,
high overhead, it "froze" in the head-back first phase of this movement.
A Plumbeous Pigeon that was, I think, in sight of another (which I was
unable to see until I later flushed it) paraded in much the same manner as the
Picazuro Pigeon (Goodwin 1964), striding to and fro on a horizontal bough
with head lowered and rump feathers erected.
106
Preening and sunbathing movements were identical to those of other
pigeons known to me. One bird, that was watched from above as it preened,
sometimes had the rump feathers fully erected but the upper tail coverts
quite flat. The uropygial gland was not thereby exposed as it is in comparable
postures of gamebirds and passerines. I have not seen any pigeon make use of
this gland when preening, as gamebirds, waterfowl and passerines so con-
spicuously do, although some have claimed that C. livia sometimes does so.
THE RUFOUS PIGEON
Observations on the male in the Belem Zoo were made for short periods
several times from 18th to 25 th November. The bird was in good condition,
very tame, and quite unafraid of being watched. The only other pigeons in the
large aviary were a number of Grey-fronted Doves. The Rufous Pigeon made
frequent sexual approaches to these but met with indifference or avoidance
from most and active aggression from one of them.
Voice: I transcribed the advertising coo as (i oo-6o, cook-cook, 66-66".
It was usually repeated from two to five times. The two shorter notes were
always interspersed by only two longer ones; thus, if given twice the phrase
would go "66-66, cook-cook, 66-66, cook-cook, 66-66". The bird in-
variably began and ended with two long notes. If allowance is made for
slight differences of interpretation this transcription agrees well enough with
those of Skutch (1964) and Belcher & Smooker (1936).
A single, purring or snoring "croorr", very similar to that of the Plumbeous
Pigeon, was sometimes given after or in intervals between uttering the ad-
vertising coo. The display coo was a single slightly purring "coor", re-
peated at each bow.
Some behaviour patterns : The bowing display was seen on only one occasion.
It consisted of a deep bow, repeated several times. The bird's head came to
the level of, or possibly a little below the rather narrow metal rod on which
it was perching. His tail went up as he bowed but both Dr. Snow and I had
the impression that this was probably a compensatory balancing movement
rather than an integral part of the display. The Rufous Pigeon was almost
sideways on to and only slightly turned towards the Grey-fronted Dove to
which he was displaying, but it is quite likely that the narrow perch prevented
a more frontal display.
What was obviously friendly nodding (Goodwin 1956b) was shown to-
wards Grey-fronted Doves whenever one of the latter came within about
four or five feet of the Rufous Pigeon. It was similar to the nodding of many
old world Columba species except that the neck seemed to be thrown more
forward at the start of the movement. At the culmination of each nod the bill
was at an angle of about 40 to 45 degrees from the substrate.
Wing ruffling occurred suddenly between bouts of nodding. As with C.
pica^uro (Goodwin 1964) and the White-crowned Pigeon C. leucocephaia, it
consisted of holding the slightly unfolded wings out from the body and
making a quick vibrating movement with them, which produced a sharp
rustling sound. This movement has also been recorded for the Spotted
Pigeon C. maculosa (Russell 191 3).
The flight-intention movements were like those of C. plumbea except for
being more leisurely. This difference was, I think, solely due to the fact that
this Rufous Pigeon was only seen to give them when he was clearly not in
the least afraid.
107
SUMMARY
Some calls and behaviour patterns of the Plumbeous and Rufous Pigeons
are described. Both utter a call which is alike in the two species and similar
to a probably homologous call of the Picazuro Pigeon. The flight intention
movements of these three species are also closely similar, possibly identical.
The Rufous Pigeon has a wing ruffling movement similar to those of the
Picazuro and White-crowned Pigeons.
References :
Belcher, C. & Smooker, G. D. 1936. Birds of the Colony of Trinidad and Tobago. Part III.
Ibis (6)13: 1-35.
Goodwin, D. 1956a. Observations on the voice and some displays of certain pigeons.
Avicult. Mag. 62: 17-23, 62-70.
— 1956b. The significance of some behaviour patterns of pigeons. Bird Study 3: 25-37.
— 1964. Some calls and displays of the Picazuro Pigeon. Condor 66: 418-422.
— 1970. Pigeons and doves of the world. 2nd ed. London: Brit. Mus. (Nat. Hist.).
Russell, W. S. H. (Marquess of Tavistock). 1913. Foreign doves at liberty. Avicult. Mag.
(5)3: 123-132.
Skutch, A. F. 1964. Life histories of Central American pigeons. Wilson Bull. 76: 211-247.
The breeding of the Madagascar Banded Kestrel
by J. F. R. Colebrook- Robjent
Received 2jth May, 1973
The Banded Kestrel Falco ^oniventris is confined to Madagascar. There are five
Falco species on the island of which three are residents ; the other two breed-
ing falcons are F. n. newtoni and F. peregrinus radama. The Banded Kestrel is
stated by Rand (1936) to be distributed from sea level to 1,000 m in the
forests of the Humid East, etc. "About Maroantsetra and Andapa (i.e. in the
north-east of Madagascar) it was fairly common, but was rare elsewhere".
Brown & Amadon (1968) summarise what little is known about this falcon,
and state under breeding habits that nothing is recorded. The following is
taken from my notes made during a recent seven-week trip to Madagascar.
However, the period covered by this paper is from 5th to 26th September
1972 when I was in the northern Humid East.
Locality, Altitude and Habitat: The area is the Manantenina River system.
From the Bekuna Stream, outside the eastern boundary of the Reserve
Naturelle de Marojejy, the valley follows the Bangwabe which runs into the
Manantenina River near Mandeny Village. The village of Manantenina is
situated where the river of that name joins the Lokoho, about three miles down
river from Mandeny. This area lies in the Prefecture of Sambava, Province
of Diego Suarez, and is 50 to 100 metres a.s.l. The valley bottom is mostly
cultivated with vanilla, coffee, rice, bananas, and coconuts, interspersed with
second-growth and strips of remnant forest. The crests of the flanking ridges
are still partially covered with degraded rain forest.
General: This falcon is usually referred to as a kestrel and its nearest
relatives are assumed to be the aberrant kestrels F. dickinsoni and F. ardosiaceus,
both of Africa (Brown & Amadon 1968). The same authorities, following
Swann (1924-1936), state that this species is uncommon. But Rand's quoted
statement more accurately reflects the status of this bird in the Manantenina
area. The Banded Kestrel was certainly "fairly common" in this valley, the
Madagascar Kestrel F. newtoni being very common. It was never seen to
108
hover (it apparently does so rarely), but neither was the more "typical"
kestrel newtoni ever seen to hover. As Rand states, it remains perched on dead
branches for long periods.
Voice: "A sharp scream and a chattering 'kek-kek-kek'" (Brown &
Amadon). In my experience it gave a low call or was often silent, but as it
often perched in trees above running water it is possible that its call was not
heard on some occasions. I was watching a pair of these falcons on 6th
September at Manantenina. They were feeding on small reptiles (probably
chameleons) on the exposed bare branches of A.lbi%pja trees. After picking
off some small prey from a branch one of these birds uttered a faint "tck-tck-
tck-tck-tck" opening its mouth wide as it did so, each note being rendered at
intervals of more than a second. I then lost sight of the bird, but could still
hear it "tcking". I eventually spotted the bird again. It had entered a thickly
foliaged large tree, still "tcking", and was making its way slowly down
towards a large basin-shaped epiphyte growing on a lateral branch. It then
dropped down into the centre of the epiphyte and briefly shuffled about
before hopping out and starting again to "tck". It then repeated the per-
formance over again. See also under Breeding.
Food: Brown & Amadon noted that the principal food items are insects
and small reptiles (including chameleons) and also some birds, the former
being taken both on the ground and in vegetation. All certainly identified
prey noted by me was chameleons taken from trees. In the crop of one speci-
men collected was a small chameleon in six pieces ; in the stomach were the
remains of another chameleon. All prey was collected from vegetation,
usually high up on the upper branches of bare Albi^ia or other leafy trees.
This falcon has acute eye-sight. After perching immobile for some time they
suddenly take off with a rapid flight and swoop up into the foliage of trees
200 or more metres away and return with a chameleon. Once a bird plunged
perhaps 100 metres into very thick-foliaged rain forest appearing seconds
later with its usual prey. Unlike most falcons, the Banded frequently enters
thick vegetation. Chameleons are placed securely in a small fork of a branch
the head being eaten first. If caught by the male the remainder of the prey is
then frequently offered to the female who eats it from the neck to the tail.
Fifteen minutes or more is taken to finish one chameleon, after which the
bird remains perched for perhaps 20 minutes with only the occasional slight
digestive head-bobbing.
Breeding: As stated earlier nothing has been recorded. F. dickinsoni breeds
usually in Boras sus and other palms, sometimes in the domed nests of the
Hamerkop Scopus umbretta, and once in a hole in an iron bridge (Brooke &
Howells 1 971). F. ardosiaceus commonly utilises Hamerkop's nests, also old
nests of other birds and holes in trees (Brown & Amadon).
Although the Hamerkop is common in Madagascar, I did not see these
birds or their conspicuous nests in the Manantenina-Marojejy area. After
witnessing the remarkable behaviour previously noted (see Voice) I sus-
pected that epiphytes (which were very common) would be used by <%oniventris
for breeding, but it was not until 24th September that this was proved to be
the case.
On that date I was investigating the nest-hole of a pp.tr of Madagascar
Kestrels in a copse of remnant (degraded) forest near the Bangwabe River.
A Banded Kestrel was seen to be perched high up on a dead tree at the edge
of forest on a nearby ridge. I had taken the precaution of posting a boy as a
look-out while I was engaged in climbing to the newtoni nest. After changing
its perch twice the boy signalled that the falcon was heading towards me. It
109
flew swiftly and low towards the copse, entering very low and suddenly
swooping up into the fork of a large and tall forest tree. It immediately
disappeared from sight in a large epiphyte. Unseen by me there was a second
bird already there and in less than a minute one of these falcons left the tree,
stooping in a low, circular dive and out of sight. Soon after, the remaining
bird (which was the female) dived off the epiphyte and swooped up to the
upper bare branches of another tree. This bird was then promptly offered a
small chameleon by the male which suddenly appeared with it. The birds
remained close together for a while when the male, without apparent
warning and with quivering wings, mounted the female for over five
seconds. During the act the female kept up a not very audible but high-
pitched "kik-kik-kik". After this the male flew off, directly over me (I was
not hidden), twisting away in characteristic rapid flight. It was seen to catch
another chameleon and settle inconspicuously in a nearby leafy tree.
Fifteen minutes later, the female returned to the epiphyte, standing in full
view in the middle of it and then slowly disappearing inside. I climbed up a
nearby tree and was able to look down into the epiphyte. The falcon then
stood up and departed revealing a fairly deep empty scrape at one side of the
bare, earthy interior.
The following day I returned to the copse. At mid-day one of the Mada-
gascar Kestrels flew to the Banded Kestrels' nest site, but was quickly chased
off by the female Banded which had been perched unseen nearby. The
Banded then settled on the epiphyte and rotated several times on the scrape.
It was also seen to peck about rather lethargically in the area of the scrape.
Most of the time it simply crouched or stood up in the interior or outside on
the leaves of the parasitic plant. About 30 minutes later the female flew off
and did not return for over an hour. One of the pair was later seen soaring
over the site, in bright sunlight, rather quickly in broad circles. This was
probably a form of display as it is quite uncharacteristic of normal behaviour.
From below the body appeared dark as did the under wing coverts; con-
trasting with the pale brown and faintly barred primaries and heavily barred
tail.
Later, in the same afternoon, a second nest site further up river was
investigated. This had been found by Michel Rakotonirina, my interpreter,
earlier that morning. This site was a broad epiphyte 13 metres up on the
trunk of a large forest tree standing almost alone in a line of trees and bushes
along a stream running into the Bangwabe. As we approached the male was
seen perched nearby on a branch of a dead tree. Although I had frequently
used a nearby track I had only once before noted a Banded Kestrel in this
area. Rakotonirina reported seeing one of these birds fly to the epiphyte and
later leave it to alight on the nearby dead tree where mating took place.
One of my Malgache guides then climbed the tree. As he was about to
reach the epiphyte the falcon rather slowly hopped out onto the first branch
and flew off without a protest. The climber reported a shallow empty scrape
in the centre of the epiphyte.
In the case of a third record, no precise nest site was located before I had
to leave the area on 26th September. On that date a male Banded Kestrel
was seen perched on the bare upper branches of an Albi^ia just above the
path leading to (and just outside) the village of Manantenina. This area is less
than 100 metres from where the incident described under Voice took place.
However, the falcons had not been seen again in the former area. There is no
true forest here, but a few large trees (mostly Albi^id) stand about the
vanilla and coffee plantations. As I watched, the male was joined by the larger
female when she was presented with a chameleon and soon after mounted
for less than five seconds. This female was later collected and proved to be in
breeding condition. The oviduct appeared to be swollen, the largest ovum
measuring 9 mm.
From the foregoing it would appear that clutches would not be completed
before the end of September or early October.
Epilogue: On 23rd October the tree at the second site was climbed by the
same Malgache guide as before. The nest then held three eggs of which two
were broken; all heavily incubated. Hery Christophe, the Forest Guard,
forwarded the remaining egg to me. It is yellowish, rather evenly speckled
and stippled with light yellowish-brown, the markings tending to be darker
brown on the broader end. The dull and yellowish markings are probably
due to the length of incubation. The egg measures 42-8 x 33-7 mm. The
average of 48 eggs of the much smaller F. newtoni is given as 35 -8 x 28-8
mm (Brown & Amadon).
Summary: Three pairs of the Madagascar Banded Kestrel Falco ^oniventris
were studied in north-east Madagascar during two weeks in September. Pre-
egg-laying behaviour and nesting sites are recorded for the first time,
together with the first known egg of this falcon.
Acknowledgements : My thanks are primarily due to Professor Charles Sibley,
of Yale University, for enabling me to visit Madagascar and for many other
kindnesses, not least for criticizing a preliminary draft of this paper. I was
helped in the field by Jali Makawa (a wonderful companion), Michel Rako-
tonirina (my interpreter) and Hery Christophe (and his two able assistants,
Robeson and Tizzy Antoin, who collected the egg of the Banded Kestrel).
Others who have helped in several ways are C. W. Benson, R. J. Dowsett,
P. Griveaud, and R. WagstafTe. J. M. Andriamampianina, Chef des Eaux et
Forets, was particularly helpful in granting permits and in many other ways.
References :
Brooke, R. K. & Howells, W. W. 1971. Falcons at Birchenough Bridge, Rhodesia. Ostrich
42: 142-143.
Brown, L. H. & Amadon, D. 1968. Eagles, hawks and falcons of the world. London: Country
Life Books.
Rand, A. L. 1936. The distribution and habits of Madagascar birds. Bull. Amer. Mus.
Nat. Hist. 72(5): 143-499.
Swann, H. K. 1924-36. Monograph of the birds of prey. London: Wheldon & Wesley.
A review of the Boat-billed Heron Cochlearius cochlearius
by Robert W. Dickerman
Received 29th March, 19J3
The relationship of the Boat-billed Heron Cochlearius cochlearius at the sub-
familial, familial and tribal levels within the order Ciconiiformes has recently
been discussed by Bock (1956), Cracraft (1967) and Dickerman (1971), but
the subspecific variation within the species has not been reviewed since the
description of C. c. panamensis Griscom (1926). During the course of investi-
gations on the possible role of birds in the ecology of arboviruses in Mexico
and Guatemala, and on the nesting biology of Cochlearius in Mexico (Dicker-
man & Juarez 1971 ; Juarez & Dickerman 1972), series of Boat-billed Herons
have been collected that, together with the limited amount of material already
available in museum collections, have permitted this review of the Mexican
and Central American populations. Measurements were recorded to the
nearest millimetre (Table).
FIGURE
Distribution of the races of Cochlearius cochlearius in Mexico and Central America. Open
squares, ^eledoni; circles, phillipsi; triangles, ridgwqyi (open triangles, localities reported in
Dickey & Van Rossem 1938); squares, panamensis; hexagon cochlearius. Localities with
vertical lines are type localities. Hatched region represents intergrade zone panamensis X
ridgwqyi.
TABLE
Measurements in millimetres of adult Boat-billed Herons from Central America, with mean
and standard deviation
Males
East Coast Mexico
West Coast Mexico S. to
Isthmus of Tehuantepec
Pacific coastal Chiapas (Mexico),
Guatemala and Honduras
East Coast Mexico
West Coast Mexico S. to
Isthmus of Tehuantepec
Pacific coastal Chiapas (Mexico),
Guatemala and Honduras
wing chord
265-296 (281-9)
SD 7-8n=28
259-273 (269-6)
SD3-3n=14
262-283 (271-5)
SD 6-6 n=16
268-281 (274-7)
SD4-5n=15
258-273 (264-6)
SD4-4n=10
255-274(268-3)
SD6-9n=14
tail
110-122(114-8)
SD3-5n=28
102-110 (106-6)
SD2-2n=14
105-114(110-1)
SD 3-3 n= 16
72-87 (79-9)
SD3-2n=29
71-83(79-4)
SD3-0n=16
71-83 (79-2)
SD3-1 n=14
Females
108-116 (111-8)
SD2-5n=16
100-107 (104-0)
SD 3-4n=10
105-116 (109-4)
SD3-0n=19
71-81 (74-7)
SD2-6n = 15
73-78 (75-4)
SD 1-9 n=10
66-78 (73-1)
SD 3-2n=14
exposed culmen
78-92 (85-2)
SD3-3n=29
78-88 (83-7)
SD2-8n=16
78-87 (83-6)
SD3-9n=14
72-84 (78-2)
SD3-9n=16
76-82(78-6)
SD l-8n=10
72-81 (76-7)
SD2-7n=14
Cochlearius cochlearius ^eledoni
Cancroma ^eledoni Ridgway (1885). Type locality, Mazatlan, State of
Sinaloa, Mexico.
Diagnosis: Pale "lavender" grey dorsally, forehead usually whitish, washed
with pale-grey or buff: wing and tail shorter than in the birds of the Gulf
coastal lowlands of Mexico.
Range: Pacific coastal areas of Mexico from Mazatlan, Sinaloa, south at
least to the Rio Papagallo, Guerrero (see Figure).
Cochlearius cochlearius phillipsi new subspecies
Type: Adult male no. 803,080 American Museum of Natural History,
collected ca. eight km east of Atasta, State of Campeche, Mexico, 13 th Feb-
ruary 1966 by Robert W. Dickerman (original field number RWD 13,677).
Testes 27 x 13 and 23 x 13 mm.
Diagnosis: Similar to ^eledoni but much larger (Table); paler on chest and
sides of neck than the population of the Pacific lowlands of Chiapas and
Central America.
Range: Gulf and Caribbean coastal zone of Mexico from La Pesca, State
of Tamaulipas, south to Belize. Probably intergrades with the next form in
the Caribbean lowlands of Guatemala from whence no material is available.
It is a pleasure to name this form in honour of Dr. Allan R. Phillips who is
so generous in sharing with others his knowledge of the Mexican avifauna.
Cochlearius cochlearius ridgivayi new subspecies
Type: Adult male no. 134, 122 Carnegie Museum, collected at Coyoles,
Department of Yoro, Honduras, on 17th June 1950 by Arthur C. Twomey
and Roland W. Hawkins (original field number 14,293). Noted as "breeding",
"TGE" (testes greatly enlarged).
Diagnosis: Similar dorsally to ^eledoni but breast a richer, deeper, vinaceous ;
much paler throughout than panamensis. Averages smaller than phillipsi but
larger than ^eledoni. The breeding birds of the Pacific coastal area of Chiapas
(and those of adjacent Guatemala), while showing the subspecific characters,
average slightly paler than Honduras specimens.
Range: Pacific coastal areas of State of Chiapas, Mexico; Guatemala; and at
least the Caribbean lowlands of Honduras. The range of ridgivayi is disjunct
from the southern part of the range of ^eledoni by approximately 450 km in
the Isthmus of Tehuantepec region. On the Caribbean lowlands there is a
sharply-stepped cline between Belize and the type locality of ridgivayi. Un-
fortunately no adult specimens are available from the intervening coastal
region of Guatemala. Locality records for El Salvador from whence no
specimens were examined are taken from Dickey & Van Rossem (1938).
This form is named with respect for Mr. Robert Ridgway whose works
form the backbone of American ornithology.
Cochlearius cochlearius panamensis
Cochlearius cochlearius panamensis Griscom (1926), type locality Corozal,
Canal Zone.
Diagnosis: Much darker, more olive above than the clear grey of the
northern races ; also more olive, less vinaceous, on sides of neck.
Range: Costa Rica: Boca Rio Matima, Limon, Boca Mala, Puntarenas
(Hellmayr & Conover 1948); intergrades with ridgivayi in Guanacaste.
Panama: throughout Republic except southwestern Darien (Wetmore 1965).
Unfortunately this form is exceedingly poorly represented in collections.
I have seen only six specimens that I consider to be adult.
Cochlearius cochlearius cochlearius
Cancroma Cochlearius Linnaeus (1766), type locality Cayenne.
Diagnosis: Adult, very much paler grey dorsally and on sides of neck,
upper breast and all of foreneck pure white. Immature, back and wing
coverts richer cinnamon than Central American forms.
113
Range: South America in suitable habitat from extreme southern Panama
south to northern Argentina.
To date I have not studied South American specimens of the Boat-billed
Heron and follow other authors in considering them all to represent the
nominate form.
AC KNOWLEDGEMENTS
The author wishes to acknowledge the authorities of Mexico, Guatemala
and Costa Rica for scientific permits to collect birds in their respective
Republics. Specimens collected in Costa Rica were taken during field exer-
cises with the Organization of Tropical Studies. Curators of the following
collections kindly loaned material or provided access to collections in their
care: American Museum of Natural History; Carnegie Museum; Museum of
Zoology, Louisiana State University; Museum of Comparative Zoology
(Harvard) ; Robert T. Moore Collection (Occidental College) ; United States
National Museum; and the Western Foundation of Vertebrate Zoology. The
research was supported in part by U.S. Public Health Service Research Grant
AI-6248 from the National Institute of Allergy and Infectious Diseases.
Dr. Kenneth C. Parkes kindly reviewed the manuscript, and his suggestions
have been incorporated in the paper.
References :
Bock, W. J. 1956. A generic review of the family Ardeidae (Aves). Amer. Mus. Novit. 1779.
Cracraft, J., 1957. On the systematics of the Boat-billed Heron. Auk 84: 529-533.
Dickerman, R. W. 1971. Systematic position of the Boat-billed Heron, in Dickerman,
R. W. & Juarez (1971).
Dickerman, R. W. & Juarez, L., C. 1971. Nesting studies of the Boat-billed Heron, Coch-
lear ius cochlear ius at San Bias, Nayarit, Mexico. Ardea 59: 1-16.
Dickey, D. R. & Van Rossem, A. J. 1938. The birds of El Salvador. Zool. Series, Field
Mus. Nat. Hist. 23.
Griscom, L. 1926. Ornithological results of the Mason-Spinden expedition to the Yucatan.
I. Introduction, birds of the mainland of eastern Yucatan. Amer. Mus. Novit. 235.
Hellmayr, C. E. & Conover, B. 1948. Catalogue of birds of the Americas. Zool. Series,
Field Mus. Nat. Hist. 8, part 1, no. 2.
Juarez, L., C. & Dickerman, R. W. 1972. Nestling development of Boat-billed Herons
{Cochlearius cochlearius) at San Bias, Nayarit, Mexico. Wilson Bull. 84: 456-468.
Linnaeus, C. 1766. Sy sterna naturae. Ed. 12(1): 233.
Ridgway, R. 1885. Description of a new species of Boat-billed Heron from Central America.
Proc. U.S. Nat. Mus. 8 : 93-99.
Wetmore, A. 1965. The birds of the Republic of Panama. I. Tinamidae (Tinamous) to
Rhynchopidae (Skimmers). Smiths. Misc. Coll. 150.
Composite nest of Short-crested Flycatcher
Myiarchus ferox
by P. J. K. Burton
Received yth May, 1973
During January to April 1972, I accompanied the British-Trans Americas
Expedition on its vehicle crossing of the Darien Gap (Eastern Panama and
North-western Colombia). On 26th March, I was at La Lomas de Rumie, a
spur of high ground jutting out into the Atrato swamp near the Rio Perancho.
Walking along an isolated rough road bordering the swamp, I observed a
Short-crested Flycatcher Myiarchus ferox carrying nest material. Through
watching the bird, I quickly located the nest which was about 12 feet above
114
the ground in a crevice of a low cliff flanking the road on one side. By scram-
bling up, I was able to inspect the nest site, and was surprised to find a
composite structure containing three nest cups. One of these held four eggs,
one held one &gg, and the third appeared to be still under construction. The
whole structure was composed largely of mammal fur, with snakeskin around
the edges of the cups. Apparently only one bird was visiting it. This was
eventually collected, and proved to be a female inbreeding condition, though
with no egg in the oviduct. The composite nest and eggs were also collected.
During a fall three of the eggs were broken. However, these, as well as the
two surviving eggs (both from the c/4) were all fresh and unincubated. Bird,
eggs and nest are now in the collections of the British Museum (Natural
History).
The genera Myiarchus and Myiodynastes are known to lay somewhat larger
clutches than the majority of tropical Tyrannidae, a fact which has been linked
with their hole-nesting habits by Lack (1968: 174). However, a composite
nest of this type must surely be exceptional ; it is unfortunate that it was not
possible in the available time to discover how many birds were involved.
Reference :
Lack, D. 1968. Ecological adaptations for breeding in birds. London: Methuen.
Re-identification of Kallus pectoralis deignani
by S. Dillon Ripley & Storrs L. Olson
Received 16th May, 1973
The description of the race Rallus pectoralis deignani (Ripley, 1970, Nat. Hist.
Bull. Siam Soc. 23: 367-368) from Celebes was of particular interest as the
species pectoralis was not previously known from that island and because
deignani was subsequently cited as providing a link between Rallus pectoralis
of Flores, New Guinea, and Australia, and R. mirificus of Luzon (Mayr, 1971,
Journ. Ornith. 112: 302-316). In our independent studies of the Rallidae, we
have had cause to re-examine the type and only specimen of deignani and fear
that we must report that it is actually an aberrant individual of Rallus
striatus — a species already known from Celebes.
As was noted in the original description, the type of deignani differs from
normal striatus in lacking the white barring in the primaries that is charac-
teristic of that species, and in this respect it does resemble pectoralis. However,
the degree of this barring is quite variable in striatus and the bars may be
broad, narrow, or sometimes reduced to disconnected spots. In one specimen
from Ceylon examined by Olson (American Museum of Natural History No.
545053), the barring was almost absent. The type of deignani, which is an
immature, thus represents an extreme in this occasional trend towards the
reduction of barring. It does preserve, however, the regularly spaced white
bars in the outer web of the outermost primary, which are typical of striatus
(in pectoralis the white markings on the outer web of the outer primary are
thin, irregularly spaced spots along the very edge). The specimen is larger
than most subspecies of pectoralis, except the nominate Australian form, and
has a stouter bill and somewhat more extensive white dorsal spotting than
pectoralis. In these respects it also agrees with striatus. Rallus pectoralis deignani
thus becomes a synonym of Rallus striatus striatus Linnaeus and the species
pectoralis must be removed from the Celebes list.
115
Migrant Perm's apivorus in the Indonesian Archipelago
by C. M. N. White
Received 4th June, 1973
Vaurie (1965 : 147-149) treats the west palearctic Perms apivorus, which win-
ters in Africa, and the east palearctic and tropical Asian P. ptilorhynchus as
separate species, thus following Stresemann (1940, Archiv. f. Naturgesch. 9:
137-193). Brown & Amadon (1968: 220-227) treat these taxa as conspecific.
The latter are followed here. Finsch (1901, Notes Ley den Mus. 22: 241) listed
an immature collected by Schadler on 8th December 1897 at Kisar, an island
east of Timor, as P. ptilorhynchus. The specimen is still in the Leiden Museum,
and Dr. G. F. Mees reports that it agrees with immatures of east palearctic
orientaiis wintering in Java. This record has been ignored in statements of the
winter range of orientaiis in recent literature, and probably since Finsch
published the original record. It extends the winter range of this form, at
least casually, far to the east, and indicates that casual migrants might even
reach north-west Australia.
Hartert (1896, Nov. Zool. 3: 177) also reported Pernis sp. from Saleyer,
south of the Celebes, a female with a moulting wing measuring 440 mm,
which he thought agreed with Malaysian ptilorhynchus. It is certainly much
too large for the Celebes species, P. celebensis. Stresemann (op. cit.) identified
it as orientaiis but otherwise the record has been ignored. These records from
Saleyer and Kisar bring migrant Eurasian honey buzzards east of Wallace's
Line, and add another species to the comparatively few palearctic migrants
east of that line (my tentative total of traced species is only 78).
Dr. G. F. Mees of the Rijksmuseum van Natuurlijke Historie, Leiden,
Netherlands, has kindly verified the details of the Kisar bird, and incidentally
pointed out the Saleyer record again. I am most grateful to him.
References :
Brown, L. & Amadon, D. 1968. Eagles, hawks and falcons of the world. London: Country Life
Books.
Vaurie, C. 1965. The birds of the palearctic fauna. Non-passeriformes. London: H. F. & G.
Witherby.
Non-passerine bird weights from Panama and Colombia
by P. J. K. Burton
Received yth May, 1973
Several recent papers (e.g. Britton 1970; Colston 1971, 1972) have supplied
weight data for a wide range of African birds, but comparatively little has
been published on the weights of neotropical birds. The best source is
probably Haverschmidt (1968), though Collins (1972) has provided some data
from Venezuela, and Snow & Snow (1963) from Trinidad. This paper
presents the weights of non-passerine birds collected during the Darien phase
of the British Trans- Americas Expedition, 197 1-2, and follows the same
format as Colston (1971, 1972). The specimens were all preserved in fluid as
anatomical material, and most are now in the collections of the British
Museum (Natural History); some were given to the Gorgas Memorial
Laboratory, Panama City. Since most of the birds have been preserved intact,
sex is given only for species showing clear cut sexual differences in external
features. A number of specimens have, however, been made into skeletons,
116
and these have been sexed by dissection; this is indicated in the text. All
weights were taken immediately after death using Pesola balances, and all are
given in grammes. Nomenclature follows Wetmore (1965, 1968); some
taxonomic problems are noted in the text. A few of the birds included here
are wintering nearctic species. In all, 163 weights of 79 species are listed.
A future paper will present similar data for the passerines collected.
Co-ordinates for the localities mentioned are as follows: C.I.M. (Military
Instruction Centre, near Pacora), 9 8' N., 79 15' W. ; Rio Espave (Bayano
Valley), 9 1 5 ' N., 78 45 ' W. ; La Palma, 8° 24' N., 78 9' W. ; Jaque, 7 29' N.,
78 i W.; Rio Jaque, 7 30' N., 78 5' W.; El Real, 8° 7' N., 77 45' W.;
Pinogana (Rio Tuira), 8° 8' N., 77 39' W.; Boca de Paya (Rio Tuira),
7 56' N., 77 32' W.; Rio Paya, approximately 7 56' N., 77 27' W.; Cerro
Pirre, 7 5 7' N., 77 45 ' W. ; Sautata (lower Atrato valley), 7 5 1 ' N., 77 8' W. ;
Cienagas de Tumarado (Atrato swamp), approximately 7 50' N., 76 5 8' W. ;
Rio Perancho (Atrato swamp), approx. 7 43' N., 77 10' W. The three last
localities are in Colombia, the remainder in Panama.
Podilymbus podiceps: 300, Rio Jaque, February.
Podiceps dominkus: 126, Rio Perancho, March.
Anhinga anhinga: $ (diss.) 1,500, Cienagas de Tumarado, March.
Butorides virescens: 205, Jaque, February.
Florida caerulea: 400, Rio Jaque, February.
Hydranassa tricolor: 335, La Palma, February.
Nyctanassa violacea: 635, 670, 680, 685, 710, La Palma, February.
Tigrisoma Hneatum: 815, Rio Paya, February; 755, imm. 725, El Real, March
S ar cor amphus papa: 3,200, Boca de Paya, February.
Coragyps atratus: 1,700, Pinogana, February.
Cathartes aura: 1,400, Rio Espave, January.
Elanoides forficatus: 420, Rio Perancho, March.
Rostrhamus sociabilis: imms. 340, 365, 370, Cienagas de Tumarado (1) and Rio Perancho
(2), March.
Buteo platypterus : imm. 350, Rio Espave, January.
Buteo magnirostris: 253, 265, 247, El Real, February/March; 242, 278, Boca de Paya,
February.
Busarellus nigricollis: 975, Rio Perancho, January.
Buteogallus urubitinga: imm. 1,200, Rio Paya, February.
Buteogallus anthracinus : 1,200, Boca de Paya, February. This bird belongs to the larger
nominate race. The race B. a. bangs'^ together with other smaller forms charac-
teristic of Pacific coastal mangroves, may be separable as a distinct species, B.
subtilis (see discussion in Wetmore 1965).
Herpetotheres cachinnans: 575, Boca de Paya, February.
Milvago chimachima: 395, Rio Perancho; 365, Sautata, March.
Daptrius americamts: 490, Boca de Paya, February; 505, Cerro Pirre, March.
Ortalis cinereiceps: 495, El Real, March. This species is treated by de Schauensee (1966)
as a race of O. garrida.
Ar amides cajanea: 440, El Real, March.
hater alius albigularis: 41, El Real, March.
Jacanajacana: 109, 133, El Real, March.
Belonopterus chilensis: <$ (diss.) 280, El Real, March.
Charadrius wilsonia: 53, 53, 53, 54, La Palma, February.
Tringa flavipes : 79, El Real, March.
Tringa solitaria: 50, Boca de Paya, February.
Actitis macularia: 32, La Palma, February.
Catoptrophorus semipalmatus: 233, 240, 282, 284, La Palma, February.
Ereunetes mauri: 21-7, 22-9, 25-0, La Palma, February.
Chlidonias niger : 37, 39, Jaque, February.
Columba nigrirostris: 236, La Palma, February.
Columbina talpacoti: 48, 50, La Palma, February; 51, El Real, February.
Columbina minuta: 30, C.I.M., January; 30, Sautata, March.
Claravis pretiosa: o* 73, Boca de Paya, February.
Leptotila verreauxi: 146, El Real, February.
Leptotila cassinii: 144, Rio Espave, January; 174, Rio Jaque, February.
117
Ama^ona ochrocephala: 405, Sautata, March.
Piqya cqyana: 102, C.I.M., January; 120, Jaque, February; 124, El Real, February.
Pi ay a minnta: 35, El Real, March.
Crotophaga major: 144, 149, Cienagas de Tumarado, March.
Crotophaga ani: 119, 130, La Palma, February; 94, 99, 109, Rio Jaque, February; 104,
118, imm. 59, El Real, March.
Crotophaga sulcirostris : 70, C.I.M., January.
Pulsatrix perspicillata: 885, Jaque, February.
Cbordeiles acutipennis: 6* 57, 6* 71, ? 57, Sautata, March.
Nyctidromus albicollis: 6* 53, ¥ 50, El Real, March.
Glaucis hirsuta: 5 -9, C.I.M., January; 5-2, 5-2, 7-1, Rio Espave, January.
Threnetes ruckeri: 6-2, 6-2, Rio Espave, January.
Phaethornis super cilio sus : 5-0, Rio Espave, January; 6-o, Jaque, February.
Phaethornis longuemareus : 2-3, Rio Espave, January.
Thalurania furcata: 4-1, Rio Espave, January.
Trogon melanurus: $ 112, Cerro Pirre, March.
Trogon viridis: $ 81, Cerro Pirre, March.
Trogon rufus: o* 57, Jaque, February.
Ceryle torquata: $ 330, $ 274, Jaque, February; $ 320, Rio Paya, February.
Chloroceryle ama^pna: $ 119, Rio Jaque, February; $ in, $ 130, o* 135, 6* 142, Boca de
Paya, February.
Chloroceryle americana: 6* 40, Cerro Pirre, March; $55, Rio Jaque, February; $$ 39, 55,
Rio Paya, February.
Chloroceryle inda: o* 60, ?? 53, 60, El Real, March
Chloroceryle aenea: $ 14-6, lower Atrato valley, March.
Electron platyrhynchum : 60, Rio Espave, January; 63, 64, 64, Cerro Pirre, March.
Baryphthengus martii: 198, Jaque, February. Included by de Schauensee (1966) under B.
ruficapillus.
Momotus momota: 108, 128, Rio Espave, January; 109, 119, 120, El Real, March.
Galbula ruficauda: <$<$ 25-1, 26-3, Rio Perancho, March. Both nominate ruficauda,
although this area is close to a region of hybridization with G. r. melanogenia.
Jacamerops aurea: $ 76, Rio Espave, January; $ 62, Cerro Pirre, March; $ 64, lower
Atrato valley, March.
Notharcus macrorhynchos: 109, Rio Espave, January; 99, 103, El Real, February/March.
Notharcus tectus: 33, Rio Espave, January.
Nystalus radiatus: 63, Rio Jaque, February.
Malacoptila panamensis: $ 46, juv. 42, Rio Espave, January; juv. 37, Boca de Paya,
February.
Nonnula frontalis: 15-2, 15-0, Rio Espave, January; 15-3, Sautata, March. Included
under iV. ruficapilla by de Schauensee (1966).
Monasa morphoeus: 93, 96, 98, 102, Rio Espave, January; 114, 122, Rio Paya, February;
105, 118, Cerro Pirre, March.
Pteroglossus torquatus: $ 196, Boca de Paya, February.
Selenidera spectabilis: $ 229, Cerro Pirre, March.
Picumnus olivaceus: 10 • 8, El Real, March.
Celeus loricatus: $ 74, Rio Perancho, March.
Centurus rubricapillus : 6* 55, $ 45, C.I.M., January.
Centurus pucherani: $53, Rio Espave, January; $ 62, Jaque, February; $ 52, Boca de
Paya, February.
Campephilns melanoleucos: $ 274, $ 181, Rio Espave, January.
References :
Britton, P. L. 1970. Some non-passerine bird weights from East Africa. Bull. Brit. Orn. CI.
90: 142-144, 152-154.
Collins, C. T. 1972. Weights of some birds of north-central Venezuela. Bull. Brit. Orn. CI.
92: 151-153.
Colston, P. R. 1 97 1. Additional non-passerine bird weights from East Africa. Bull. Brit.
Orn. CI. 91 : 1,10-111.
— 1972. African passerine bird weights. Bull. Brit. Orn. CI. 92: 11 5-1 16.
Haverschmidt, F. 1968. Birds of Surinam. Edinburgh & London: Oliver & Boyd.
Meyer de Schauensee, R. 1966. The Species of Birds of South America. Wynnewood, Pa:
Livingston.
Snow, D. W. & Snow, B. K. 1963. Weights and wing-lengths of some Trinidad birds.
Zoologica 48: 1 - 1 2 .
Wetmore, A. 1965. Part 1 & 1968. Part 2. The Birds of the Republic of Panama. Smiths.
Misc. Coll. 150 (1 & 2).
118
Once more : the identity and authorship of
Treron griseicauda
by G. F. Mees
Received 4th June, 1973
Sims & Warren (1955) drew renewed attention to the unsettled authorship
and identity of Treron griseicauda, which by previous authors had been either
used for the Javanese subspecies of Treron pompadora as T. p. griseicauda
G. R. Gray, 1 8 5 6, or for the Celebesian subspecies as T. p. griseicauda Wallace,
1863. Sims & Warren (195 5) discovered that Bonaparte (1854: 10) had listed
T. griseicauda as a synonym under the name T. aromatica, although his des-
cription was, in fact, based on specimens of the Javanese subspecies, and not
on T.p. aromatica from Boeroe. They concluded that T. griseicauda Bonaparte,
1854, notwithstanding the fact that it was first published as a synonym, is an
available name, and therefore proposed to call the Javanese subspecies T. p.
griseicauda Bonaparte, 1854.
A year later, Mayr (1956) objected to the treatment of Sims & Warren,
observing that according to the principles laid down at the Zoological
Congress, Copenhagen, an otherwise unavailable name does not acquire
availability by being cited subjectively in the synonymy of another name.
Subsequently, however, the following emendment of the International
Code was adopted by the XVIth International Congress of Zoology (Wash-
ington 1963): "A name first published as a synonym is not thereby made
available unless prior to 1961 it has been treated as an available name with its
original date and authorship, and either adopted as the name of a taxon or
used as a senior homonym" (Art. 1 i[d]).
There cannot be the slightest doubt that Sims & Warren, well before 1961,
have used T. griseicauda Bonaparte exactly in the meaning of Art. 1 1 (d) and
therefore have validated it. The correct name of the subspecies of Treron
pompadora occurring on Java is therefore Treron pompadora griseicauda
Bonaparte, 1854.
If it was the change in redaction of Art. 11(d) alone that affected the nomen-
clature, I might have left the matter as it was, but there are two reasons why
a renewed discussion appeared desirable. The first is that Goodwin (1967:
308), in a much-used work, has accepted Bonaparte's name, but without
explanation.
The second reason is that even if one rejects T. griseicauda Bonaparte, the
name T. griseicauda Wallace (1863a) for a bird from the Celebes is of doubtful
validity, as long as the exact date of publication of T. griseicauda Schlegel
(1863b) is not known. Schlegel (1863a) mentioned that the first nine sheets of
the "Nederlandsch Tijdschrift voor de Dierkunde" were printed around the
middle of 1862, but that: "the publication of this work can only begin with
1863". Wallace's name was published in April 1863 (cf. Duncan 1937),
Schlegel's paper was not only reviewed in the Ibis of July 1863 (p. 359-360),
but in the same number Wallace (1863b) devoted a special article to a dis-
cussion of it, which definitely shows that it must have been received some
time before July, and may well antedate Wallace's paper of April 1863.
Sanft (i960, note 12) has quoted evidence that the first part of the "Neder-
landsch Tijdschrift voor de Dierkunde" was published in 1862, as Schlegel
(1862), in a work dated August 1862, and certainly published before the end
of that year, gave full references, with page- and plate-numbers, to it {Buceros
119
pulchrirostris and Buceros nagtglasii). However, this was explained by Schlegel
(1863a) as quoted above, and later Schlegel (1873) gave 1863 as year of
publication of the pigeons described in the first part of the "Nederlandsch
Tijdschrift voor de Dierkunde". A small consequence for the nomenclature
is that not, as Peters (1945) and Sanft (i960) did, the "Nederlandsch Tijd-
schrift voor de Dierkunde" has to be cited as containing the original
descriptions of Buceros pulchrirostris and Buceros nagtglasii, but the "Museum
des Pays-Bas".
I take this opportunity to mention that if one follows Goodwin (1967) in
merging the genus Megaloprepia with Ptilinopus (personally I am unconvinced
of the necessity to do so), the species listed as Ptilinopus for mosus (Gray) by
Goodwin, must be known as Ptilinopus bernsteinii Schlegel, as correctly
pointed out by Schlegel (1863a); see also Wallace (1865 : 388). Indeed, under
the present Code (Art. 59[b]), Ptilinopus bernsteinii has been the valid name
for the species right from the moment of its publication.
References :
Bonaparte, C. L. 1854. Conspectus Generum Avium 2. Lugduni Batavorum.
Duncan, F. M. 1937. On the dates of publication of the Society's "Proceedings", 1859—
1926 (etc.). Proc. Zool. Soc. Lond. 107A: 71-84.
Goodwin, D. 1967. Pigeons and doves of the world. London: Brit. Mus. (Nat. Hist.).
Mayr, E. 1956. The names Treron griseicauda and Treron pulverulenta. Bull. Brit. Orn. CI. 76:
62.
Peters, J. L. 1945. Check-list of birds of the world 5. Cambridge, Mass.: Harv. Univ. Press.
Sanft, K. i960. Bucerotidae. Das Tierreich 76: 1-174.
Schlegel, H. 1862. Buceros. Mus. Hist. Nat. Pays-Bas 1 (mon. 1): 1-22.
— 1863a. Letter to the editor. Ibis 5 : 11 9-1 20.
— 1863b. Observations sur les colombars, voisins des Treron aromatica et vernans. Ned.
Tijdschr. Dierk. 1 : 63-71.
— 1873. Aves Columbae. Mus. Hist. Nat. Pays-Bas 4 (mon. 35): 1-180.
Sims, R. W. & Warren, R. M. L. 1955. The names of the races of the Pompadour Pigeon,
Treron pompadora (Gmelin), in Java and Celebes. Bull. Brit. Orn. CI. 75 : 96-97.
Wallace, A. R. 1863a. List of birds of the Sula Islands (east of Celebes), with descriptions
of the new species. Proc. Zool. Soc. Lond. for 1862: 333-346.
— 1863b. Notes on the fruit-pigeons of the genus Treron. Ibis 5 : 318-320.
— 1865. On the pigeons of the Malay Archipelago. Ibis (n.s.) 1 : 365-400.
St. Vincent Parrot : plumage polymorphism,
juvenile plumage and nidification
by Holly A. J. Nichols & Thomas Duncan Nichols
Received 21st April, 1973
INTRODUCTION
We have been able to re-evaluate variation in plumage colour of the St.
Vincent Parrot Ama^ona guildingii as a result of our observation of captive
and museum specimens and the first captive breeding of this species. One of
us (T.D.N.) observed plumage variation in wild parrots and some details of
nidification in a preliminary field trip (19-23 April, 1973) to St. Vincent.
PLUMAGE VARIATION
In 1967 the government of St. Vincent granted special permission to
Young Island Hotel of St. Vincent to donate two juvenile A. guildingii to
U.S. zoos. The hotel donated one to the National Zoological Park, Washing-
ton, D.C., and the second to the Houston Zoological Gardens. Since A.
guildingii is an IUCN endangered species, these two zoos and the New York
Zoological Park (Bronx Zoo) and the Chicago Zoological Park (Brookfield
Zoo), each having one specimen, instituted a co-operative breeding pro-
gramme, bringing the parrots together at the Houston Zoo. The Chicago
parrot died in 1970; its skin is now at the Louisiana State University Museum
of Zoology. The Houston Zoo purchased a fifth, apparently very old A.
guildingii in 1971. We have thus been able to closely study these five adults.
One of us was able to carefully examine an additional twenty-two captive
specimens on St. Vincent.
One of us (H.A.J.N.), a former employee of the Houston Zoo, was
responsible for the first Houston parrot, the New York parrot and the one
offspring they successfully raised in 1972. This was the first time this species
had been raised in captivity, allowing us the opportunity to study the juvenile
plumage.
The plumage coloration of A. guildingii is complex. Clark (1905) and Ridg-
way (19 1 6) have described it in detail. They describe some specimens which
are generally more green than brown, their most conspicuous difference
being primaries with the basal portion green rather than yellow. Clark and
Ridgway described these green parrots as immature, although Ridgway
cautioned in a footnote, "I strongly suspect that in reality [this plumage]
represents a dichromatic phase".
James Bond described (pers. comm.) an A. guildingii nestling he observed
in 1927 as "covered with short, dull green feathers". Allen (1961) reported
"young entirely green" probably on the basis of Bond's observation. John
Warfield, who was responsible for the two juvenile specimens the Young
Island Hotel donated to the U.S. 200s, has written (pers. comm.) that one
of the two was always more green. The second, however, he described as
being distinctly more brown. Both of these parrots are now at the Houston
Zoo and six years old. The first bird's plumage is that described by Clark
and Ridgway as immature. The second bird (the breeding female) and the
other three specimens including the juvenile parrot, now at the Houston
Zoo, have the yellow-brown plumage described by Clark and Ridgway as
adult. The juvenile plumage coloration of the one parrot raised in 1972 was,
except as noted below, virtually identical to that of the yellow-brown morph
parents. There are thus two morphs, green and yellow-brown, and the
juvenile plumage coloration of any particular bird is quite similar if not
identical to that of its eventual adult plumage. We are aware of eight green
morph and forty-eight yellow-brown morph specimens in fourteen museum
collections and ten green morph and seventeen yellow-brown morph
specimens in captivity.
Clark noted that of the eight specimens he collected, no two were exactly
alike in plumage. Similarly we must say it is difficult to find any specimens
of the five at the Houston Zoo, of the twenty-two observed in captivity on
St. Vincent, and the fifty-six museum skins with like plumage. Although
the differences between green and yellow-brown morph specimens are dis-
tinct, many yellow-brown morph specimens, such as the mounted specimen
in the City of Liverpool Museums, have some areas of green on the back or
carpalia. Some birds, such as a yellow-brown morph in the possession of
William Miller of St. Vincent, appear to have a more yellowish tone to the
breast and fewer greenish areas (we have not been able to positively classify
this variation).
There is also considerable variation in other plumage characteristics such
as the width of the central blue-green tail band, head coloration, neck
coloration, and amount of black feather edging. LSUMZ 69139 for example
is unusual in having a pale yellow chin and adjoining malar region, and an
orange throat and posterior malar region, the usual blue being mostly
confined to the nape and green to the hind neck. An apparent partial lutino,
owned by Miller, has many deep yellow feathers on the nape and hind neck.
Two specimens, BM 89.1.20.307 and AMNH 417,403, have collars with very
heavy black edging.
Although the excessive variation of head coloration may eventually be
understood in terms of distinct morphs, we would only point out the existence
of two extremes and their approximate occurence. The breeding male at the
Houston Zoo, which is typical of one extreme, has his eye almost surrounded
with feathers which are at least tipped with blue-violet, the forehead and
anterior crown only being white. The Houston breeding female is an example
of the other extreme. There is much more white on her head, the eye always
having been completely surrounded by white. The lores, forehead, crown,
and anterior portion of the auriculars are generally white except for some pale
yellow in the posterior crown. We have not been able to classify the varia-
tions between these two extremes, there being several patterns of cartenoid
and melanin deposition resulting in heads with mixtures of white, yellow,
orange-brown and blue-violet. The head coloration of the Houston juvenile
parrot for example is similar to the breeding female's except for some yellow
and blue posterior to the eye. We are aware of nine specimens of the first
extreme, seven of the second and forty-two we are not able to classify. All
variations in head coloration occur in both green and yellow-brown morph
parrots.
We have not been able to detect any changes in head coloration from photo-
graphs taken over several years of the parrots now at the Houston Zoo. The
breeding male for example looked the same in 1972 as when Arthur W.
Ambler photographed it in 1959 (Allen op. cit. 100). (In the last few months,
however, diet changes of the five parrots at the Houston Zoo have been
responsible for eliminating much yellow from their head plumage). Many of
the individual parrots observed on St. Vincent in the collections of Miller
and John Houser were known to be four of five years old. As a group these
birds displayed a wide variety of stable plumage characteristics, such as
referred to in this paper, specifically having all varieties of head coloration.
(We do not believe that any of the plumage colour variation of the twenty-
two captive specimens observed on St. Vincent resulted from dietary
abnormalities; they were all well fed.) We conclude age does not normally
affect head plumage coloration.
We have not detected correlation between sex and any of the plumage
characteristics.
FIELD OBSERVATIONS
As Clark indicated, the distinction between the green and yellow-brown
morphs (which he considered young and adult) is relatively easy to make in
the field. I (T.D.N.) observed both morphs in the upper Buccament Valley.
Although I was not able to accurately count the number of each morph
observed, I. A. Earle Kirby, his associate, and I observed or heard approxi-
mately twenty-five parrots in nine hours. The parrots were eating the abund-
ant fruit of Pouteria multi flora.
We found two nest sites, one at 1,000 ft. in an Ormosia monosperma, about
40 ft. up the trunk, and one at 1,500 ft. in a Dacryodes exce/sa, about 80 ft. up
the trunk. D. excelsa is usually hollow and I was told that it was the normal
nesting tree.
I
Kirby's associate, an unusually observant native naturalist, said that he
had never seen "a clutch larger than the normal two". A clutch of two would
be in agreement with the 1905 report of Clark, the 1927 observation by Bond
(pers. comm.), the 1930 report of Porter, and the clutch of two eggs laid in
1972 at the Houston Zoo.
One egg shell was obtained (laid by a captive bird) from Warfield as a
donation to Louisiana State University Museum of Zoology. It measures
38-1 by 30-1, as compared to 43 -8 mm by 32-6 mm for the first (infertile)
of the two eggs laid at Houston in 1972, and 46 -6 mm by 38 -8 mm for the
egg reported by Harrison & Holyoak (1970).
SUMMARY
Ama^ona guildingii has been found to display a wide variety of plumage
polymorphism. Two major morphs are identified, which had previously been
considered as the immature and adult plumages. The juvenile plumage of
any individual is the same as its adult plumage. A few details of nidification,
and observations made on St. Vincent are reported.
AC KNOWLEDGEMENTS
We thank the following persons for describing or lending specimens of
the indicated institutions: Dr. A. Schwartz (Miami), Dr. R. A. Paynter Jr.
(Museum of Comparative Zoology), Dr. E. R. Blake (Field Museum of
Natural History), Dr. J. Bond and Dr. J. V. Griswold (Academy of Natural
Sciences), Dr. J. M. Forshaw and Dr. J. P. O'Neill (Louisiana State Uni-
versity Museum of Zoology), Dr. W. B. King and Dr. R. L. Zusi (Smith-
sonian Institution), Dr. M. Gochfeld and Mr. D. Ewert (American Museum
of Natural History), Dr. K. C. Parkes (Carnegie Museum), Dr. J. C. Barlow
and Dr. R. D. James (Royal Ontario Museum), Dr. P. J. Morgan (City of
Liverpool Museums), Dr. E. H. Stickney (Yale Peabody Museum of Natural
History), Dr. D. D. Lonsdale (Bermuda Aquarium, Museum and Zoo),
Dr. D. W. Snow (British Museum, Natural History), and Mr. C. W. Benson
(Cambridge University Museum of Zoology). We thank Dr. W. G. Conway,
Dr. J. Bell and Mr. A. Ambler for providing information and photographs
of specimens previously at the New York Zoological Park, Mr. R. Small for
photographs of the specimen previously at the Chicago Zoological Park,
and Mr. J. W. Houser, Mr. J. Warfield, Mr. W. Miller and Dr. I. A. E.
Kirby for their assistance on St. Vincent.
References :
Allen, R. P. 1961. Birds of the Caribbean. 223. New York: Viking.
Clark, A. H. 1905. Lesser Antillean birds (Ama^pna guildingii). Proc. Bost. Soc. N. H. 32:
267-271.
Harrison, C. J. O. & Holyoak, D. T. 1970. Apparently undescribed parrot eggs in the col-
lection of the British Museum (Natural History). Bull. Brit. Orn. CI. 90: 42-46.
Porter, S. 1930. Notes on the rare parrots of the genus Amazona. Avic. Mag. (4)8: 2-12.
Ridgway, R. 1916. Birds of North and Middle America. Part VII: 225-227. Washington:
Govt. Print. Off.
The humerus of Ichthyornis as a taxonomically isolating
character
by C. J. 0. Harrison
Received 1 6th June, 19J3
Two small sea-birds of the genus Ichthyornis were described by Marsh (1880)
from the Cretaceous of Kansas. According to him they possessed true teeth,
resembling in this respect Hesperornis, with which they co-existed. A number
123
of skeletal elements were found and Marsh commented "In their powerful
wings and small legs and feet they remind one of the terns, and, according
to present evidence, they were aquatic birds of similar life and habits". He
used the skeleton of the Royal Tern as a basis for his reconstruction of /.
dispar. Gregory (1952) suggested that the toothed jaws associated with the
species were those of reptiles and that Ichthyornis species had bills like those of
modern birds. Following from this, with Marsh's comments on their tern-
like appearance well-established in literature, there was a tendency to suggest
Ichthyornis as the probable ancestor of the modern Charadriiformes (Fisher
1967, Brodkorb 1971).
More recently Walker (1967) and Gingerish (1972) have argued that
Ichthyornis species were toothed, and have found additional material which
appears to confirm this, but quite apart from the question of the presence or
absence of teeth Ichthyornis species also show at least one other morphological
peculiarity, which was remarked on by Marsh but which has received little
attention since. This is the size and position of the crests on the humerus.
Marsh (pp. cit.) comments on the exceptional size and position of the deltoid
crest in his description of the humerus ; but in discussing the reconstruction
of Ichthyornis he states that "the locomotive organs of Ichthyornis are so
similar to those of typical birds that they present no such interesting mechani-
cal problems as were suggested by the skeleton of Hesperornis" . The position
of the deltoid crest is, however, quite exceptional.
Normally the deltoid crest is an elongated ridge along the external side of
the shaft of the humerus at the proximal end. It provides, inter a/ia, an attach-
ment surface for the large pectoralis major muscle which creates the main
Figure 1. Proximal, palmar and external views of the proximal ends of left humeri of
(a) Ichthyornis dispar and (b) Platalea leucorodia.
power for the downstroke of the wing in flight. It is usually an elongated
prominent flange of bone, stout at the base and thinner towards the edge;
but the degree of development varies in different species and in some it is
little more than a stout ridge. In all modern birds it projects palmarly from
124
the external palmar edge of the shaft and is usually almost at right-angles to
the transversely-elongated head (Fig. ib). It may curve laterally the further
it projects from the shaft, and towards the edge in some species the outer
portion may be at an angle of c. 50-60 degrees to the head.
In Ichthyornis dispar, the species for which complete humeri are known, the
deltoid crest is large and long and projects laterally, continuing the line of
the transverse proximal head (Fig. ia). It forms a thin flange almost at right-
angles to the position of those on modern birds. It could be suggested that
specimens had undergone some crushing and distortion, but I have examined
a specimen of a proximal end of a humerus completely prepared from its
matrix, and on this, although the hollow shaft has been crushed, the surface
of the outer part of the crest and the region adjacent to the proximal head
show no evidence of the displacement needed to change the position of this
flange had it originally been in a position comparable with that of a modern
bird.
The structural modification was not a general early avian feature. As
Yalden (1972) has pointed out, the deltoid crest of Archaeopteryx h extensively
developed, but an examination of the specimen of the latter in the British
Museum (Natural History) reveals that in this bird the deltoid crest projects
in a similar palmar plane to those of modern birds. The humerus of Hes-
perornis is reduced to a slender strip of bone, and the only humerus showing
some similarity to that of Ichthyornis is one from the Lower Cretaceous of the
Weald, currently being described (Harrison & Walker, in press), on which,
although the outer portion of the deltoid crest is broken away, the portion
present appears to show similar structure to that of Ichthyornis and may be
earlier evidence of this modification.
Another peculiarity of the humerus of Ichthyornis is the apparent absence of
the bicipital crest. This normally forms a rounded, laterally-projecting crest
on the internal side of the shaft adjacent to the proximal head, bordering a
smooth surface which projects a little palmarly. The size of the bicipital
surface and crest varies from one species to another in modern birds, but
these are never wholly absent except on the greatly reduced humeri of some
ratites.
With the humerus in position on the body of a modern bird the deltoid
crest projects horizontally and slightly ventrally along the upper edge of the
humerus, but if the humerus of Ichthyornis is placed in a similar position the
crest projects dorsally. There do not appear to be modifications of the
articulating surfaces of the humerus of the latter which would indicate some
other position for this bone. Marsh's reconstruction of /. dispar (1880)
shows this dorsal projection of the deltoid crest.
The position of the deltoid crest in modern birds is presumably that which
provides the most efficient surface angle for the functioning of the flight
muscles. For that of Ichthyornis to work in comparable fashion would involve
considerable forward rotation of the humerus and hence of the whole wing.
It is improbable that Ichthyornis would show evolutionary divergence to
produce a less efficient system and much more likely that the modification of
the humerus indicates a specialised muscle system which probably involved
modification of the mode of flight. Since the changes shown involve an
absence of, or change in angle of, surfaces used for the attachment of major
muscles for wing movement it is likely that the mode of flight was peculiar
and may have involved more gliding, although the modifications of humeri in
modern gliding birds such as the Procellariiformes are quite unlike those of
Ichthyornis.
125
Assuming that the arrangement of crest on the proximal end of the humerus
in Archaeopteryx and in modern birds represents the normal condition in the
main evolutionary stem in birds, then the development of the deltoid crest
and loss of the bicipital crest in Ichthyornis represent a marked evolutionary
divergence from this, not paralleled in modern forms. It is unlikely that an
evolving Ichthyornis phylogenetic line would regain the more typical humeral
structures, and in this character, as in the probable possession of teeth, it
appears to have been an evolutionary blind end, having no ancestral affinities
to any known subsequent avian taxa.
References :
Brodkorb, P. 1971. Origin and evolution of birds, in D. S. Farner & J. R. King, eds.
Avian Biology 1 : 20-55. New York: Academic Press.
Fisher, J. 1967. Fossil birds and their adaptive radiation, in The Fossil record: 133-154, 75 3—
762. London: Geol. Soc.
Gingerish, F. D. 1972. A new partial mandible of Ichthyornis. Condor 74: 471-472.
Gregory, J. T. 1952. The jaws of the Cretaceous toothed birds Ichthyornis and Hesperornis.
Condor 54: 73-88.
Harrison, C. J. O. & Walker, C. A. in press. Wyleyia, a new bird humerus from the Lower
Cretaceous of England. Palaeontology 16.
Marsh, O. C. 1880. Odontornithes ; a monograph on the extinct toothed birds of North America.
Washington : Govt. Printing Office.
Walker, M. V. 1967. Revival of interest in the toothed birds of Kansas. Trans. Kansas
Acad. Sci. 70: 60-66.
Yalden, D. W. 1971. The flying ability of Archaeopteryx. Ibis 113: 349-356.
The relative numbers of migrant and resident rollers
in eastern Kenya
by L. H. & B. E. Brown
Received 22nd May, 1973
In February 1968, when touring the Tsavo National Park of eastern Kenya
with members of the Fauna Preservation Society, it was noted that the
European Roller Cor arias garrulus was very numerous. The average density
was estimated at 2-3 per mile (1 -2 — 1 -cj/km) of road traversed, which, on
the basis of the mean visibility on either side, gave a density of 60-90 rollers
per square mile (23-3 5 /km 2 ). Possibly rollers may have concentrated along
roads though there is no direct evidence that they were not evenly spread
through the habitat. If similar densities prevailed throughout the Tsavo
Park there would have been 480-720,000 in the park itself, while the total
in the thornbush area of eastern Kenya alone would be of the order of 2-3
million. This may appear a huge figure; but doubtless these birds collect here
from an enormous area of European and Asian breeding habitat.
In the eastern Kenya thornbush there are two main rainy seasons, the
long rains extending in good years from late March to mid-May and in bad
from mid- April to mid-May; and the short rains, extending from late October
to late December in a good and early November to early December in a bad. i
As a rule the April-May rains are the heaviest, and April the peak rainfall j
month. However, in some areas the November-December rains are less
erratic. The dry seasons which follow, January-March and June-September, !
also differ, that between January and March being shorter but hotter than j
the mid year dry season, when July and August are usually cool and overcast. I
126
European Rollers are present in Kenya from October to late April,
straggling into May; but are only numerous from early November to March,
that is, in the short rains and succeeding dry season. They usually arrive
in numbers when the first heavy storms of the short rains occur in early
November and bring out an abundance of insect life. Other migrant in-
sectivorous species, for instance Hobbies Falco subbuteo, are also very common
at this time. Migrant European Rollers then greatly outnumber the local
resident species, the Lilac-breasted Roller Coracias caudata and the Rufous-
crowned Roller Coracias naevia. This is a feature common to other migrant
genera, for instance hirundines and wagtails.
In an attempt to obtain more exact quantitative data on numbers of
resident and migrant species counts were made in 1969-70 of all rollers
along stretches of the main Nairobi-Mombasa road at different times of year
on journeys to and from the coast. The counts were not made at "ideal"
times but fitted in with other affairs. Some of them have already been briefly
referred to by Moreau (1972: 181), but are given here in greater detail and
related to season and ecological conditions, in Tables I— III.
TABLE I
Early long rains counts, March-April
Food abundant by April. European Rollers on northward' passage
Date
Conditions
Sector
C. garrulus
C. caudata
C. naevia
Total
31.3.69
Early rain. Heavy
passage N. of
C. garrulus
I
II
III
2
103
68
9
5
2
4
7
2
15
115
72
Total
173
16
13
202
31.3.70
Poor early rains;
bush very dry,
few rollers
I
II
III
8
2
2
2
2
10
2
Total
8
2
4
14
25.4.69
Wet, after
I
main departure
II
10
8
18
of C. garrulus
III
18
16
21
55
Total
18
26
29
73
16.4.70
Wet, after
I
1
1
early light
storms
II
III
7
1
2
4
11
1
20
6
Total
8
6
13
27
Total March counts
181
18
17
216
Total April counts
26
32
42
100
Total March-April
207
50
59
316
Ratio C. garrulus: other spp. March 4-9:1
April 0-35 : 1
127
TABLE II
Dry season, August-September
European Rollers absent. Food scarce
Date
Conditions
Sector
C. garrulus C. caudata C. naevia Total
5.8.69
Dry.
I
j
O
!
Cool
II
O
O
O
III
O
O
O
30.8.69
Dry.
I
O
O
O
Sunny
II
2
O
2
III
O
O
O
Total o 3 03
22.8.70 & So few of any species seen that counts not considered worth analysing.
5.9.70 Even local species very scarce.
TABLE III
End short rains, December-January
Food sometimes abundant. European Rollers present
Date
Conditions
Sector
C. garrulus
C. caudata
C. naevia
Total
20.12.69
Lush after
good rains
I
II
III
1
79
58
1
5
1
2
19
1
4
103
60
Total
167
4.1.70
Dry: vegetation
I
5
1
6
still long, but
II
78
2
11
9 1
leaf fallen
III
117
2
119
Total 200
Total Dec. /Jan. counts 338
216
383
Total, all counts
545
65
92
702
Ratio: C. garrulus: other spp. Dec. /Jan. 7-5:1
Ratio: C. garrulus: other spp. year round 3-5:1
The road was divided, mainly for convenience, but also because of
ecological differences, into three sectors : — I, Kiboko-Mtito Andei (48 miles,
77km); II, Mtito-Andei-Voi (61 miles, 98 km); and III, Voi-Mackinnon
Road (46 miles, 74 km). Originally, all were Acacia-Commiphora-Adansonia
woodland with a decreasing cline eastward from about 750 to 500 mm of
annual rainfall, Kiboko to Mackinnon Road. However, Sector I, the wettest,
supports a fairly heavy human population and consequently a low population
of elephants Loxodonta africana^ with the result that much of the woodland,
including large baobab trees Adansonia, survives. Sector II passes almost
entirely through the Tsavo National Park, where an enormous elephant
population has practically destroyed the woodland including almost all I
Commiphora and Adansonia; and Sector III skirts the boundary of the National !
Park (strictly respected by the resident elephants) so that both sides of the
road are again wooded. This sector is also somewhat drier than the others,
with more stunted Commiphora woodland. Certain other birds, for instance I
128
the migrant Grasshopper Buzzard Butastur rufipennis, are common here in
November-December and not elsewhere.
The resident species C. caudata and C. naevia occur the year round, but
their numbers also fluctuate, indicating local movements. Slender evidence
suggests that their main breeding season may be the long rains, April-May,
so they should reach peak numbers in May-June. Insects and other suitable
prey are then quite as, if not more abundant than they are in November-
December, when C. garrulus is also present in far greater numbers and could
compete with the residents for such food. However, C. naevia at least, which
is the commonest resident roller in Commiphora woodland (C. caudata prefers
moister areas), also becomes quite numerous in November-December and
breeds in both main rainy seasons.
From the counts, however, it appears that the numbers of both species of
resident rollers increase sharply between March and April, at a time when
European Rollers are departing northwards and the first storms of the long
rains have fallen. Their numbers approximately double between late March
and late April, presumably preparatory to breeding. These rollers too are
clearly nomadic or migratory, for, after breeding in the long rains, they also
become very scarce in the latter part of the June-October dry season. C.
naevia^ which is the commoner species and better adapted to Commiphora
woodland, apparently also becomes common again in the short rains
November-December.
So far the evidence does not support any suggestion that a heavy popu-
lation of elephants in Sector II has depressed the breeding population of
resident species by depriving them of breeding sites. The total of all counts
of resident species in Sector II, the worst affected by elephants is 84, cf. 5 2
for Sector III, and 21 for Sector 1, the woodland least affected, with abundant
breeding holes in baobabs. Corrected for distance, these figures become 84,
71 and 22, still suggesting that the highest resident roller populations are in
areas worst damaged by elephants.
Possibly the large numbers of dead but not yet completely rotted trees
and stumps resulting from elephant damage may actually have provided more
breeding holes than formerly existed, so favouring the resident rollers, of
which C. naevia certainly also breeds in banks of streams. However, this
suggestion requires further investigation.
In general it seems probable that:
(1) The total numbers of any species of roller inhabiting this thornbush
is not necessarily limited by food supply. The numbers of resident rollers
living in the area in April and May, when near their peak, are always very
much lower than the combined total of all species present, or of European
Rollers alone, from November to March. The total of all European Rollers
in four December-March counts is 519 (mean 130/count), despite one very
low count of eight on 31.3.70, after most of the birds had left. Without this
low count the average of the other three counts would have been 170, which
may be compared with 26 (mean 13) in two April counts. In other words
although food, which would be relatively scarce at the end of the January-
March dry season, becomes abundant again in April, the total numbers of
rollers, especially of European Rollers, decrease. Those of both resident
species in the same counts are 80 in December-March (mean 27) cf. 74 (mean
37) in April. Year round there are on average 3 -5 European Rollers for one
of any resident species while in December-March the proportion is 6 • 5 : 1
in favour of the European Roller. This is so despite the fact that C. naevia at
129
least is a larger and more powerful species than C. garrulus, killing inter alia
snakes up to at least 50 cm long.
(ii) That the numbers of all species fluctuate greatly from year to year
according to the rains and that rollers in general are more nomadic in the
African tropics than is generally realised. Local movements of the Abyssinian
Roller C. abyssinica have been related to the dry and wet seasons by Brown
(1946: 194), and Elgood et al. (1973: 35) find all Coracias and Eurystomus
rollers migratory in Nigeria. In Kenya and Ethiopia the Broad-billed Roller
Eurystomus glaucurus is notably nomadic, coming into breeding condition and
display at once with the onset of rains in, for instance, western Ethiopia.
(Brown unpub.) The Somalian-Ethiopian race of the Lilac-breasted Roller,
C. c. lorti, is certainly migratory, arriving in Ethiopia-Somalia in the early
rains in March, breeding and then departing south to spend the off season in
northern Kenya (Archer & Godman 1961 : 704; Urban & Brown: 1971 64).
Regular counts en route to the coast were discontinued after 1970, but it was
noted that the numbers of European Rollers fell greatly in 1971-72, a drought
year, though increased again in 1972-73, when the short rains were again
good (102 seen on Sector II and III on 8.1.73).
(iii) The numbers of resident rollers may be limited by territorial behaviour
rather than food supply. In April there should be strong competition for
available nesting holes in trees and in banks (sometimes used by C. naevid)
between rollers, glossy starlings, hornbills, wood-hoopoes and others.
However, there is little actual evidence that shortage of holes reduces the
population, rather the reverse. The European Roller in. winter, on the other
hand, merely has to find a suitable perch from which it can forage in a tem-
porary feeding area, using trees, telegraph wires, termite hills and any dead
stump of the type recently abundantly provided by the Tsavo elephants. It
may thus be able to exploit the sometimes over-abundant available food supply
in the November-March period more effectively than any breeding resident
species, whose territorial requirements may be evolved to survive in dry
years when food is relatively very short.
References :
Archer, G. & Godman, E. M. 1961. The Birds of British Somaliland and the Gulf of Aden III.
Edinburgh & London: Oliver & Boyd.
Brown, L. H. 1946. Birds and I. London: Michael Joseph.
Elgood, J. H., Fry, C. H. & Dowsett, R. J. 1973. African migrants in Nigeria (first part).
Ibis 115 : 1-45.
Moreau, R. E. 1972. The Palaearctic- African bird migration systems. London & New York:
Academic Press.
Urban, E. K. & Brown, L. H. 1971. A checklist of the birds of Ethiopia. Addis Ababa:
Haile Sellassie I University.
The voice of Sarothrura insularis with further notes
on members of the genus
by Stuart Keith
Received pth June, 1973
In April, May and June, 1971, 1 travelled extensively in Madagascar in com-
pany with Mr. A. D. Forbes-Watson and Mr. D. A. Turner. Having pre-
viously recorded the voices of six of the African species of Sarothrura, I was
130
most anxious to hear and record the voice of Sarothrura insularis ^ endemic to
Madagascar. The only information I had on its call was the description by
Rand (1936: 360), and I speculated that this call might resemble one of the
calls of Sarothura rufa (Keith, Benson & Irwin 1970: 65).
I guessed wrong. We were accompanied on part of our journey by M.
Georges Randrianosolo from O.R.S.T.O.M. in Tananarive, and when he
identified a mysterious call in the forest of Tsarafidy as that of Sarothrura
insularis^ I frankly didn't believe him — it didn't remotely resemble any sound
I had ever heard from a Sarothrura. He was later proved right, however, when
calling birds were seen and recorded near Didy, 2 5th-29th April, 1971.
The vocalisation I would like to name the "full call" lasts about 20 seconds
and may be described as follows: first one or two short trilled notes are
given, followed by several loud double notes, which are followed in turn by
a long series of loud single notes of similar quality. These single notes
gradually accelerate and at the same time decrease slightly in pitch, becoming
quite faint toward the end. This is a loud, high-pitched, and ringing call
capable of carrying for nearly half a mile. It may be written: "drr - drr -
KEEKEE - KEEKEE - KEEKEE - KEE - KEE - KEE - KEE - KEEK
- KEEK - KICK - KICK - kick - kick - kick - kick - kikikikikikikiki
". This vocalisation I take to be territorial in nature and it might there-
fore be better described as a song (for the distinction between "song" and
"call", see discussion in Keith et al. y ioc. cit.: 15).
Other vocalisations of S. insularis are mostly either shortened versions of,
or excerpts from, this full call. The introductory trill is frequently dispensed
with, and sometimes the double notes are too. Sometimes there is just a short
burst of notes lasting 5-10 seconds, while at other times the full call may be
considerably lengthened, the bird continuing to give a series of low "kiks",
unevenly spaced. This last was usually given in response to song playback,
or to the call of a nearby rival, and I take it to be a grumbling "annoyance
call" similar to that noted for S. boehmi {loc. cit.: 56).
Sonagrams of four vocalisations of S. insularis are shown in Fig. i-
Sonagrams A and B are both taken from a full call ; A shows the double note
followed by two single notes, while in B there is just a series of single notes.
Notice that in B the notes are spaced more closely together — this is from the
latter part of the call where the notes are speeded up. In A a distant bird can
faintly be seen calling "over" the near one — only the upper harmonic has
registered. Sonagram C shows the beginning of a short, loud call which only
lasted about 10 seconds — notice there are no introductory notes, and that
it very quickly accelerates and descends in pitch. Sonagram D shows a
different vocalisation, not part of a full call and a "call" rather than a "song".
It has a dry, slightly trilled quality and the rhythm is irregular — notice the
space between the notes increases in this particular excerpt.
Of course at this stage, with limited experience, I cannot ascribe "meaning"
to any of these vocalisations, except to repeat that the full call seems fairly
clearly to be the territorial call or "song". At Didy the birds were very vocal —
when one bird called it would frequently set off several others in the neigh-
bourhood. I cannot say whether they were breeding or not, since we neither
collected specimens nor found nests. They may not have been — this was
April, and Rand {loc. cit.) only records breeding in September and October.
However, the birds might be expected to breed during the rainy season,
which lasts from November to April, so they may have been at the tail end
of their breeding season during our visit (see following page).
131
Song playback readily elicited a response, often from several birds. Some
of these were far enough apart to have been males on separate territories,
while others were close enough to have been two birds of a pair. Calling by
a suspected pair was similarly noted for S. lugens (Keith et al. 1970: 50) in
response to song playback (see also Keith et. al., loc. cit.: 13-15 for discus-
sion of female rallid calls).
o
g*S
,. *•«
-.,,«.i <*,(
ti^MiMtt^m^m^mmmmmmtm^m^mmmm
1 4 ' 1.6 ' 1.8 : I l.O ' ll.2 ' ll.4 ' Ut^hf^i&O^Z^^lztr
>- ' 1.2
O fij
O
_J
I |i | ii|^ U |M i |M| .i , l i 2 | ,i i IU .i | {i r V ^ 'lJ ' »C ' y"|ii1 i' i
^ ^ MR* ,«*•< fW«:* jSM*M f m*%U 0W***
<T L2 1 I4 1 IS 1 IJEp ILCT IJ2 1 II4 1 Its' ilB 1 I2D 1 I2T 124^
** • Ai
f^i
1.2 ' 14 1.6 ' IS Tt.0 11.2 ' 1.4 ' 1 1.6 ' TiS I2.O' 12.2 124
TIME IN SECONDS
Figure i . Sonagr ams of vocalisations of Sarothrura insular is. In order from top to bottom
A, B, C, D (see text).
We may now compare the voice of Sarothrura insularis with the voices of j
other members of the genus. The voices of ayresii and watersi ate still un- I
known, but those of the remaining six are depicted in the sonagrams in
Keith et al. (loc. cit.: 1 2-1 5). It can be seen at once that insularis is unlike any
of the others, and this is even more obvious to the human ear. The African
birds have low, moaning songs and fairly low, hollow calls. The songs are
132
all at iooo cycles or lower, while calls are mostly pitched around 2000 cycles,
with overtones up to 3000 cycles. The calls of insular is are pitched at about
4000 cycles, with an undertone at 2000. The first notes in Sonagram C are at
5000 cycles, while the upper reaches in C are at 6000. To the human ear this
difference sounds even greater than the sonagrams would suggest; insularis
sounds high and ringing, the African birds low and hollow. Besides the
difference in pitch, the form of the full call of insularis is also unique — no
African bird has an accelerating, descending call.
In view of these great differences it is impossible to say to which African
species insularis is most closely related on the basis of voice. We speculated
(Joe. cit.: 71) that on morphological grounds affinis and insularis seem so
closely related as to form a superspecies. This supposed closeness is not
backed up by the vocalisations, and I would prefer now to abandon this
thought. Affinis has a low moaning song, and in fact the call of lugens seems
the closest to that of insularis , although it is still very different. The vocali-
sations of insularis cannot be used as a clue to its relationships with other
members of the genus ; furthermore, their difference is now such that I would
hesitate to class insularis as a member of any superspecies.
Further notes on Sarothrura insularis
I must here report a locality error and discuss its consequences. In our
species account of Sarothrura insularis (Keith, Benson & Irwin 1970: 64-65)
we frequently comment on the occurrence and breeding of the species at
Manombo "in the subdesert". This was particularly interesting as the only
evidence for the bird's occurring in this habitat.
In 1 97 1 Messrs. Forbes- Watson, Turner and I discovered that there are
two towns named Manombo in Madagascar. The principal one is located at
22 56' S., 43 29' E., on the southwest coast north of Tulear, in the sub-
desert. Benson, Irwin and I assumed that this was the place indicated.
However, there is also a tiny settlement named Manombo at 32 10' S.,
47 50' E., on the souths// coast south of Farafangana. Whereas it is evident
from the route map in Rand (1936: 147) that both Manombos were visited by
his expedition, the locality he gives (Joe. cit.: 360) for the two nests he dis-
covered is "Manombo (southeast)" (italics mine). They are further described
as being "in a grassy field near the forest". This is the normal habitat for the
species, and all our remarks based on its occurrence at Manombo in the
subdesert (a most unlikely habitat) must now be revised.
Rand (Joe. cit.: 360) gives the habitat of the species as "the secondary
brush, the grassland on the edge of the forest, and the little clearings in the
forest". We can now add marshes to this list of habitats. At our campsite a
few km north of Didy (about 3 5 km south-southeast of Ambatondrazaka),
while some birds inhabited the forest edge, a larger number were living in
the open marsh nearby. This is an extensive flat area of long grass, reeds and
sedge inhabited by such species as Rallus madagascariensis, Dryolimnas cuvieri
and Calamocichla newtoni. The birds were frequently seen scuttling across the
track which crosses the marsh, and many more called from the marsh than
from the woods. We did however also see an adult male and a fully-grown
young male, the latter with a rufous wash already appearing on the forehead
and chin, walking on the forest floor a few hundred yards from the marsh.
Like many species in Madagascar, Sarothrura insularis is clearly tolerant of a
variety of habitats. This further confirms my supposition (see above) that the
birds had about finished breeding at the time of our visit.
133
Notes on other members of the genus
Sarothrura elegans
An adult male was collected at Koobi Fora on Lake Rudolf, Kenya,
(3 55' N., 36 10' E.) by A. Duff-McKay and R. E. Leakey on 26th April,
1 97 1. Details were given by Duff-McKay to A. D. Forbes-Watson in Nairobi
and then sent to me. Koobi Fora is a very dry locality with almost no ground
cover — scrubby bushes and small trees. Near the lake shore, where the bird
was collected, there is a sparse covering of "ouch ! grass", and about one
km from the collecting point there are extensive reed beds on the shore of
the lake. When collected the bird was hiding in the thatched roof of a small
hut, where it had apparently been seen on the two previous days. Duff-
McKay further reports that Mrs. R. E. Leakey had seen "some" a week or so
before running around in the "ouch ! grass".
This is a most interesting record. The normal range of the species in
eastern Africa is from Cape Province, South Africa north to central Kenya
and southern Sudan. There are only four records from north and east of this
range: a female from Somalia (the type of "S. buryz"); a male from south-
central Ethiopia (Roux & Benson 1969), and two aural records: Benson
(1947: 49) heard the call near Mega, southern Ethiopia, and I taped the song
of a bird at Mt. Marsabit, northern Kenya. Koobi Fora, on the shore of
Lake Rudolf at its northeastern end, is about 1 60 miles northwest of Mars-
abit and 140 miles west of Mega, these being the two nearest recorded
localities. While perhaps not strictly being a range extension, Koobi Fora
being south of a line drawn between the localities in Somalia and southern
Sudan where the bird has been taken, this certainly adds an interesting locality
to its sparsely documented range in northeastern Africa, and is only the
third specimen for the area, and the second male.
Perhaps of even greater interest is the question, what were the birds doing
there? This is a most unusual habitat for the species. Whereas S. elegans is
tolerant of a variety of habitats, including forest, scrub, long grass and bush
near forest, and even overgrown cultivation, it is always associated with
thick, often impenetrable vegetation. It is therefore most surprising to find
birds running around in dry grass in semidesert country. Benson's bird from
Mega was in juniper woods, the bird from near Shashamane, south-central
Ethiopia, was taken in the undergrowth of a degraded Podocarpus forest, and
my bird at Marsabit was in montane forest — all within the range of typical
habitats. The only bird taken in habitat which sounds similar to that at Lake
Rudolf is the unique specimen from Somalia, which according to Archer &
Godman (1937: 334) was collected at the foot of the Wagar Mountains at
700 ft. around hot springs in desolate country. Is it possible that there is a
population of S. elegans in northeastern Africa which has adapted to living in
waterside habitats in semi-arid country? Only further field work will tell.
The condition of the gonads of the Koobi Fora bird was not noted, but
elsewhere in Kenya males in breeding condition have been taken in February,
April and May, so the presence of birds at Lake Rudolf in April might indi-
cate breeding.
The only other possibility is that the birds were on migration. There are
two objections to this idea, however. First, S. elegans is not known to be a
migrant elsewhere in its range, at least not a long-distance migrant. A few
occurrences suggest local movement (summarised in Keith et al. 1 970 : 3 3-34)
but most birds are sedentary and many probably live in the same locality
year-round, in common with almost all African forest birds. Secondly,
!34
supposing these birds were migrants, where could they have been migrating
from and to? Why would birds leave their forest habitat in southern Ethiopia,
southern Sudan, or northern Kenya and cross hundreds of miles of desert to
reach a different country? There is no reason to suppose that any of the above
places becomes inhospitable at any time of year, nor do the birds have any
rallid competitors in these places. Until further evidence is forthcoming, the
status of the birds at Lake Rudolf must remain a mystery.
I compared the Rudolf specimen with birds in the American Museum of
Natural History from southern Africa belonging to the nominate race and it
agrees with them on colour. With a wing of 95 mm, however, it is the longest
winged specimen on record. We list (loc. cit. : 3 7) the wing measurements of
all specimens we could find, and the wing range of males is 80-94, females
84-93 mm. Its other measurements (culmen 16, tarsus 25, middle toe 30 mm)
fall within the ranges for males. The long wing might perhaps indicate a
disposition to migrate.
My associates Benson, Irwin and I are indebted to Mr. Terence Oatley for
drawing our attention to two points arising from our paper (Keith et al.
1 970). On p. 40 we mention the nest of Sarothrura rufa found by Pooley (1962:
45) on the Usutu River in Zululand. Oatley writes (in litt. to Irwin) "from
the habitat and calls described by Pooley this was a genuine Sarothrura all
right but almost certainly elegans. The greenish tinged eggs were probably
white but so described because of the green cast of the light in the dense
forest understorey".
Secondly, Oatley notes a point of geographical synonymy on p. 33. Under
"Movements" we cite two records by Astley Maberly, one from "the foot-
hills of the northern Drakensburg" and the second from "the northeastern
Transvaal". These, Oatley tells us, are one and the same place.
We are likewise indebted to the Rev. Dr. William Serle for pointing out
to us (in litt. to Benson) that the precise type locality of Sarothrura elegans
reichenovi (Sharpe) is known to be Buea, ex-British Cameroons. We had
stated (p. 36) that the precise type locality was not known, but Serle drew
our attention to a paper by Reichenow (1892) dealing with a collection of
birds made Dr. Preuss in the Cameroons, from which Sharpe's type was
taken. S. elegans appears on p. 178, and it is evident that the collecting locality
is Buea at 950 m altitude.
Sarothrura pulchra
With regard to the call of this species, Serle (in litt. to Benson) says that on two
occasions his African skinner Gilbert shot female birds while calling. The call
of the female "resembled that of the male but was not nearly so loud, was
higher pitched, and the notes followed each other so closely that it was
almost like a trill and the whole performance occupied a shorter time than
the usual male sextuplicated call". Gilbert also believes that the female also
has a call just like that of the male. This is most welcome supporting evidence
for the view that female rails call as well as males.
It is now known that Sarothrura is one of those rallid genera in which the
primaries are moulted alternatively rather than simultaneously (vide e.g.
Stresemann & Stresemann 1966). Since, however, there are not very many
collected specimens of Sarothrura in moult, it may be worth mentioning a
further specimen. Mortensen (1971), writing on a collection of birds from
Liberia and Guinea, describes (p. 1 16) a male S. pulchra with one primary half
grown which was shot on a nest with two eggs.
i35
Acknowledgements
I would like to thank my co-authors of the Sarothrura monograph, Messrs.
C. W. Benson and M. P. Stuart Irwin, for many helpful suggestions, and the
three of us are grateful to Mr. T. Oatley and the Rev. Dr. W. Serle for allow-
ing the information they supplied us with to be included in this paper. I
would also like to acknowledge the help of my colleagues Messrs. Forbes-
Watson and Turner in tracking down and identifying the voices of Sarothrura
insularis, and the three of us are likewise grateful to M. Georges Randriano-
solo for pointing out the call of the bird and helping us find it. We would
also like to thank M. Roederer, Director of O.R.S.T.O.M., for permitting
M. Randrianosolo to accompany us in Madagascar.
References :
Archer, G. & Godman, E. M. 1937. The birds of British Somaliland and the Gulf of Aden 2.
London: Gurney & Jackson.
Benson, C. W. 1947. Notes on the birds of southern Abyssinia. Ibis 89: 29-50.
Keith, Stuart, Benson, C. W. & Irwin, M. P. Stuart. 1970. The genus Sarothrura (Aves,
Rallidae). Bull. Amer. Mus. Nat. Hist. 143(1).
Mortensen, P. 1971. A collection of birds from Liberia and Guinea (Aves). Steenstrupia
(Zool. Mus. Copenhagen) 1: 115-125.
Pooley, A. C. 1962. Red-chested Pigmy Rail Sarothrura rufa. Lammergier 2(2): 45.
Rand, A. L. 1936. The distribution and habits of Madagascar birds. Bull. Amer. Mus. Nat.
Hist. 72(5): 143-499-
Reichenow, A. 1892. Zur vogelfauna von Kamerun. Journ. Ornith. 40: 177-195.
Roux, F. & Benson, C. W. 1969. The Buff-spotted Flufftail Sarothrura elegans in Ethiopia.
Bull. Brit. Orn. CI. 89: 119- 120.
Stresemann, E. & Stresemann, V. 1966. Die mauser der vogel. Journ. Ornith. 107, suppl.
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Published by the BRITISH ORNITHOLOGISTS' CLUB and printed by
The Caxton and Holmesdale Press, 104 London Road, Sevenoaks, Kent.
Bulletin of the
British Ornithologists' Club
Edited by
C. W. BENSON
Volume 93 No. 4
December 1973
Committee
Sir Hugh Elliott, Bt., o.b.e. {Chairman)
J. H. Elgood {Vice-Chairman)
C. W. Benson, o.b.e. {Editor)
R. E. F. Peal {Hon. Secretary)
P. Tate {Hon. Treasurer)
J. H. R. Boswall
Dr. P. J. K. Burton
P. Hogg
C. J. Mead
Bulletin of the
BRITISH ORNITHOLOGISTS' CLUB
l
Vol. 93 No. 4 Published: 20 December, 1973
The six hundred and eighty-fourth meeting of the Club was held at the
Cafe Royal, 68 Regent Street, London, W.i, on Tuesday, i8th September
1973 at 7 p.m.
Chairman: Sir Hugh Elliott, Bt., o.b.e., present 23 members and 22 guests
The speaker was Dr. Leslie H. Brown, o.b.e., b.sc, who addressed the
Club on British Birds of Prey. The following is a precis of his address : —
Despite legislative and educational measures of recent decades, our diurnal
raptor fauna still strikes a visitor from Africa as impoverished and threatened.
Of the 25 species, nine are vagrants, of which three (Sea Eagle, Black Kite,
Red-footed Falcon) could conceivably some day stay to breed. Another two
are boreal migrants, which although fairly regular are unlikely to do so
(Gyrfalcon, Rough-legged Buzzard). Of the remaining 14 breeding species,
four are represented by under 10 pairs (Honey Buzzard, Marsh Harrier,
Osprey, Goshawk — the first two limited by ecological factors). There are
10-50 pairs of Red Kite and Montagu's Harrier, 50-100 of Hobby, and 100-
500 of Hen Harrier, Golden Eagle, Merlin and Peregrine. This leaves only
three species with populations exceeding 500 pairs and reasonably common,
especially in the west where agricultural pesticides and gamekeepers, mainly
pesticides, but even more so gamekeepers, are less prevalently hostile to
Kestrel, Sparrowhawk and Buzzard respectively.
Notwithstanding some successes (Hen Harrier, Osprey, Red Kite) — none
of them unmarred by human interference — the conservation picture is black.
Thus, since i960, the Golden Eagle has continued its decline: now, out of
about 280 pairs, 220 breed but 80-90 of them fail through persecution by
gamekeepers and shepherds, casual disturbance, egg-collecting and other
causes, in descending order of importance. The proportional loss is far
higher — reducing breeding success, for example, from 0-83 to 0-56 per pair
per annum — than that resulting from the much publicized eagle-shoots in
America, which accounted at most for 2 • 5 % of a population conservatively
estimated at 50-100,000.
The speaker emphasized the view that gamekeeping (for which land-
owners should bear a full share of responsibility) remains the major threat,
though other factors discussed were poisons and pesticides, disturbance
(the disappearance of 5-16 pairs of eagles in Deeside in the last 12 years was
attributed to this cause), habitat destruction, ecological limitations, and
"unknown" (Merlin). It followed that, while pesticide monitoring and re-
search, behavioural research (of which the needs and opportunities were far
from fully appreciated and exploited, particularly in the case of such species
as Merlin and such subjects as food and breeding success, even of Buzzard
and Kestrel), extension of reserves, and possibly reintroductions, are all
necessary and desirable, their success depends on a much more effective
reformation of attitudes and above all law-enforcement than anything yet
achieved.
i37
The juvenile plumage of Pseudhirundo griseopyga and
identity of "Vom Swallows"
by C. H. Fry
Received ijth August, 197)
In his paper on the African swallow Pseudhirundo (Hirundo) griseopyga (Sunde-
vall), Dowsett (1972) referred to two peculiar hirundines netted and released
alive by Dr. J. R. Lang at Vom (c^o'N., 8°5o'E.) in central Nigeria on 20th
March 1968. Colour photographs and measurements showed them to
resemble P. griseopyga, a small swallow with dark brown crown, glossy blue-
black back, and greyish rump, but there were some distinctive plumage
differences. The opinion had previously gained wide currency that they
might represent a new species, although Dowsett thought they would prove
to be some form of griseopyga.
On 26th April 1973 Dr. Lang netted three further "Vom swallows" and he
is certain that they are the same as the 1968 birds. I have examined the three
in the flesh at the British Museum (Natural History) and there is no doubt
that they are juvenile P. griseopyga, vindicating Dowsett's judgement. The
uncertainty arose in the first place because the juvenile plumage, which
differs markedly from the adult, is not at all adequately described in any of
the standard African texts, despite there being already a few juvenile skins at
the Museum. A full description is being published in the Bulletin of the
Nigerian Ornithologists' Society, and the salient details are as follows: —
Mantle, scapulars and flight feathers brown, entirely lacking the adults'
strong blue gloss (two specimens) or with blue gloss discernible only on
close inspection (one specimen); rump feathers grey-brown with 2 mm wide
pale fawn tips; marked superciliary stripe formed by narrow (< imm)
whitish tips and edges to feathers at sides of crown, the stripe broadening
behind the eye and even extending onto the nape and in strong contrast with
the mask formed by black-brown lores and ear coverts; pale buffy tips to
secondaries, inner primaries and greater wing coverts; tail notch about 25
mm deep (up to 59 mm in adult).
Thus in the field juvenile P. griseopyga will somewhat resemble a fork-
tailed Riparia: a small hirundine with mid brown upperparts with paler
rump, and black mask contrasting with white superciliary stripe and white
underparts.
At close range the juvenile appears scaly, owing to the more or less narrow
buffy edges to its brown feathers. This applies also to adults in fresh plumage,
since at all moults newly-grown feathers are narrowly buff edged. The pale
tips probably abrade away rapidly. In addition both juveniles and adults have
a salmon-pink wash on the freshly-grown white feathers of the chin, throat
and breast. This colour also seems to disappear soon; a Nigerian nestling
near to fledging on 9th May (B.M. reg. no. 1955.59.159) is pink below,
while neither the Vom specimens nor one of the same age and season (9th
April) from further north in Nigeria show any pink (B.M. reg. no. 1933.8.2.
27.1). Adult P. g. melbina from Liberia and P. g. gertrudis from Nigeria and
Cameroun have warm buff- or salmon-pink chins and throats when freshly
moulted in August, but September and later skins are without any pink.
The residential status of P. griseopyga in Nigeria has been discussed by
Elgood, Fry & Dowsett (1973) and little can be added on the strength of the
138
Vom records. If it breeds on the Jos Plateau, on which Vom is situated at
1,310 m, it must do so extremely sparsely since in 14 years Dr. Lang has seen
none other than those netted. It may be significant that, as Dr. Lang reports,
there was a small migratory movement at Vom when the 1973 birds were
netted, with Palaearctic passerines and the first plateau record of Turnix nana
on the same day. The swallows, which I consider had fledged within four
weeks, may have been migrants dispersing northward from Niger-Benue
breeding grounds (Bannerman 1953).
References :
Bannerman, D. A. 1953. The Birds of West and Equatorial Africa. 2 vols. Edinburgh
& London: Oliver & Boyd.
Dowsett, R. J. 1972. Geographical variation in Pseudhirundo griseopyga. Bull. Brit. Orn. CI.
92: 97-100.
Elgood, J. H., Fry, C. H. & Dowsett, R. J. 1973. African migrants in Nigeria. Ibis 115:
i-45, 375-4H-
A new race of Thekla Lark in Harrar, Ethiopia
by C. Erard & G. Jarry
Received 21st August, 197 3
In classifying the specimens recently brought back from Ethiopia by various
expeditions from the Laboratoire de Zoologie (Mammiferes et Oiseaux) du
Museum National d'Histoire Naturelle de Paris, we noticed three Thekla
Larks from Harrar which at once seemed to us peculiar. These three speci-
mens have been compared by one of us (C.E.) with the extensive material in
the British Museum (Natural History). We thank Dr. D. W. Snow and
I. C. J. Galbraith for facilities provided and C. W. Benson who translated
our paper. These birds seem to us sufficiently homogeneous and distinct
from other races of Galerida theklae to require their own name. Accordingly
we propose:
Galerida theklae harrarensis, subsp. no v.
Description: This new form has no close resemblance to the populations
of the high plateaus of Ethiopia attached to praetermissa (Blanford), the
i colour of which is very dark, especially the reddish fawn underparts. It is
akin to ellioti Hartert of Somalia, from which it is nevertheless easily dis-
tinguishable. Harrarensis is in effect darker, less sandy: the streaks on the
back are clearly of a brown which is more blackish in tone, these dark
centres to the feathers being wider and much better defined. Below, the
pectoral streaks are heavier. Finally, an important character, the bill of
harrarensis is shorter than that of ellioti, and it is also clearly finer.
These specimens have also been compared with G. t. huriensis Benson of
; northern Kenya. Six specimens of the latter in worn plumage, comparable in
this respect to our specimens, are very easily distinguished by their darker
colour. The mantle of harrarensis is less densely streaked, the dark centres of
the feathers being narrower. The feathers of the rump and upper-tail coverts
; are of an almost uniform pinkish brown contrasting with the rest of the
upperparts, whereas in huriensis the rump and upper-tail coverts are merely
fringed with a brown of a reddish rather than a pinkish tone and the centre
139
is dark brown. Below, huriensis is more streaked on the chest, the markings
being particularly large and densely distributed. Furthermore, huriensis has
the bill longer and stouter, and has much less light fawn on the outer
rectrices.
Distribution: At present harrarensis is only known from three specimens
collected in the north of the province of Harrar on 18th March 1971:^, 5 km
west of Jigjigga (9°2i'N, 42°46'E), altitude 1,700 m; go, 32 km south-east
of Jigjigga (9°i4'N, 43°o2'E), altitude 1,550 m. The first one was obtained
in a steppe of low acacia bushes scattered over stony ground; the other two
in an analogous habitat but much more rocky owing to the presence of the
bed of a dry wadi. At these two localities, and in between, this form was
relatively common. It did not appear to be breeding at this time.
Type: <$ adult, 5 km west of Jigjigga, 18th March 1971. Collectors' no:
2,794. In Museum National d'Histoire Naturelle, Paris.
Measurements of type: Wing 101, tail 57, bill (from skull) 14-5, hind-claw
11 mm.
Measurements of material as a whole: The following is a summary of measure-
ments, of material all of the same degree of wear of plumage:
Wing
Tail Bill from
skull
G. t. harrarensis
Hind-claw
*o*o*
100, 101
55, 57 14-5 (both)
10-5, 11
9
93
52 14-5
G. t. ellioti
11
7 0*0*
99-103 • 5
51-60 14-5-17-5
8-12
av.
ioi-8
56-5 15-7
G. t. huriensis
9.9
5 0*0*
97-101
51-5-60-5 15-17
11-5-15
av.
99-1
56-0 16-1
12-2
The type of harrarensis weighed 30 g; the male from 32 km south-east of
Jigjigga 32 g, the female therefrom 3ig.
Remarks: Urban & Brown (1971: 71) record only G. t. praetermissa and
huriensis from Ethiopia. The latter should strictly speaking only figure in an
inventory of the birds of Ethiopia as "probable". It has never been collected
or recorded within Ethiopia, although Benson (1945) obtained his specimens
only five miles south of the frontier with Kenya but never met with the
species during his important investigations in Sidamo. Urban & Brown do
not record G. t. ellioti. In this respect they follow White (1961: 50), who
records this race only from the interior of British Somaliland south to
Galkayu and Merca in Italian Somalia. Now, the type-locality of ellioti is
Daghabur (8°n'N, 43°3i / E), which is in the Ethiopian part of the Ogaden,
that is to say, in the south of the province of Harrar. Thus it is right to admit
this form to the list of the birds of Ethiopia and to investigate its exact
distribution in the Ogaden.
References :
Benson, C. W. 1945. Observations from Kenya Colony, Uganda and Tanganyika Territory.
Ibis 87: 90-95.
Urban, E. K. & Brown, L. H. 1971. A check-list of the birds of Ethiopia. Addis Ababa:
Haile Sellassie I University Press.
White, C. M. N. 1961. A revised check list of African broadbilts, pittas, larks, swallows, wagtails
and pipits. Lusaka: Government Printer.
140
A new race of Thekla Lark in Bale, Ethiopia
by C. Erard <& R. de Naurois
Received 21st August, 19J3
A study by one of us (R.N.), on the evolution of larks of the genus Galerida,
led us to consider jointly the populations of G. theklae of the high plateaus of
Ethiopia, until now considered as constituting the subspecies praetermissa
(Blanford). Accordingly we have examined the material brought back by
recent expeditions to Ethiopia organised by the Laboratoire de Zoologie
(Mammiferes et Oiseaux) du Museum National d'Histoire Naturelle de Paris
and that housed in the collections of the British Museum (Natural History).
It is apparent to us that the birds of Bale are distinguishable from those from
other high plateaus of Ethiopia, from which they are separated by the Rift
Valley. They seem to constitute a discrete form, which we propose to call,
in memory of our friend the late Francois Hue:
Galerida theklae hue/, subsp. nov.
Description: This form has no close resemblance to G. t. ellioti Hartert or
G t. harrarensis Erard & Jarry (see immediately above), which are light
coloured with a pronounced sandy tone. It is however near to G. t. praeter-
missa and huriensis Benson. In comparison with these two races, huei has the
streaking of the upperparts distinctly blacker and more intense (in worn
plumage, the back and the top of the head appear almost uniformly black).
The pale borders of the feathers of the mantle are narrower and lighter, more
creamy or very light buff, less pinkish brown, this being particularly clear on
the neck, where there is the outline of a pale band. The cheeks are likewise
more intensely streaked than in praetermissa. On the underparts huei is white,
very slightly washed with light buff as in huriensis, whereas praetermissa is a
dark pinky brown, only rarely pale in colour. The streaking of the chest, of a
much less heavy aspect than in huriensis, is similar to that of praetermissa, but
the streaks are heavier and more densely distributed. The throat is white as
in huriensis but much less streaked ; in praetermissa the throat is buffy. The
beak of huei is intermediate between that of praetermissa (long and slender)
and that of huriensis (long and stout).
Distribution: G. t. huei inhabits the mountains of Bale where it has been
collected at Dinsho (7°o6'N, 39°47'E) at an altitude of 3,200 m, and where
the species is relatively common, inhabiting steppes and plains of a medium
altitude (cf. Dorst & Roux 1972: 208).
We likewise attribute to huei, although not identical in all characters, two
specimens collected on 28th February 1971 at Asella (7°5 8'N, 39°o4'E), in
Arussi, at an altitude of 2,640 m, on the edge of degraded forest intersected
by cultivation. They are in fresh plumage, with a very dark pigmentation,
although the edges of the feathers of the mantle are more like those in
praetermissa (that is, more pinkish brown) of the Addis Ababa plateau. On
the other hand, the underparts are much as in huei. The measurements of
these two specimens are included below with those of that form.
It is probably also huei which Dorst & Roux (1973) found well represented
at Koffole (7°o5'N, 38°45'E), in Arussi, on arid steppes at an altitude of
2,550 m.
Type: g adult, Dinsho, 14th March 1968, specimen no. 24 collected by
party composed of Prof. J. Dorst, F. Roux and J. Chauvency. In Museum
National d'Histoire Naturelle, Paris.
141
Measurements and weight of type: Wing 103, tail 54-5, bill (from skull) 17-5,
hind-claw 1 i • . 5 mm. Weight 34 g.
Measurements and weights of material as a rvhole: We compare below only
praetermissa and huei; for data for huriensis, el/ioti and harrarensis, the other
races recorded from Ethiopia, see Erard & Jarry above. The three items
indicate respectively mean, range and number of specimens examined.
Wing
Tail
Bill from skull
Hind-claw
Weight
milli
metres
grammes
G.
/. praetermissa
101 --]
97-5 - 107
21
55-7
5 49-5 - 60
20
16-6
i5'5 - i7'5
20
i3'5
11 — 16-5
19
35-2
32- 39
14'
95-3
91 -99
5
51-2
48 - 54
5
16-1
15 - i7-5
5
G. t. huei
13-1
11 - 15-5
5
34-0
30- 36
3
103-7
101 - 107
4
96-5
95-5-97
3
54-6
52-5 - 57-5
4
51-0
50- 52-5
3
17-0
16 - 18
4
15
1
13-5
11-5 - 15
4
14-3
12 - 16- 5
3
35-0
34- 37
4
33'3
32- 34
3
??
99
Remarks: The existence of this dark form huei was foreseen by Friedmann
(1937: 31), who remarked that the birds collected by the Childs Frick
Expedition on the plateau of Arussi were blacker on the back and head than
those of Shoa and that they "may represent an undescribed form". Friedmann
refrained from naming it, not knowing the extent of individual variation in
praetermissa; this all the more because Blanford (1870: 381) had named
"A.. (G.) arenicola? Tristram, var. fusca" a very dark specimen obtained at
Ashangi (i2°3 5'N, 39°39'E). It seems to us that praetermissa does show a
certain variation in intensity of pigmentation. But in spite of an examination
of the long series in the British Museum we have not found in this form any
individual as dark and showing the characters such as we have given in the
definition of huei.
Three specimens from the vicinity of Ambo (8°57'N, 37°58'E) collected
in October 1971 between 2,200 and 2,400 m in altitude show a slight ap-
proach to huei. However, a careful examination shows that they are pale
below, the pectoral striation is of the praetermissa type, being narrower and
more diffuse than in huei, the throat is washed with light burly, the upperparts
are certainly dark but not to the extent found in huei, while the pale margins
to the feathers of the mantle do not accord with this latter but rather with
those in praetermissa.
References :
Blanford, W. T. 1870. Observations on the geology and apology of Abyssinia. London: Macmillan.
Dorst, J. & Roux, F. 1972. Esquisse ecologique sur l'avifaune des monts du Bale, Ethiopie.
L'Oiseau et R.F.O. 42: 203-240.
— 1973. L'avifaune des forets de Podocarpus de la province de l'Arussi, Ethiopie. L'Oiseau
et R.F.O. 43: in press.
Friedmann, H. 1937. Birds collected by the Childs Frick Expedition to Ethiopia and Kenya
Colony. Part 2 — passeres. U.S. Nat. Mus. Bull. 153: 506 pp.
142
A new race of Pectoral-patch Cisticola from Cameroun
by C. Chappuis <& C. Erard
Received 21st August, 1975
During a scientific journey which took us from Tchad to Gabon, we had the
opportunity in October 1972 to traverse the montane area of Adamaoua in
central Cameroun. On 17th October, in the area of the Mbangi mountains
(see Plate 90 in Bartholomew 1956), at 40 km north of Ngaoundere (ca.
7°45'N, i3°2 5'E) and altitude 1,400 m, we discovered a population of the
Pectoral-patch Cisticola Cisticola brunnescens Heuglin on a plateau covered
with meadows consisting of very short grass on stony ground, and strewn
with strands of small area of bushes, relatively little crowded, meagre and
low (height o-8o to 1 m). This bushy vegetation was more luxurious in
hollows and drainage lines, developing into shrubs (height 2 to 3 m).
The Pectoral-patch Cisticolas were frequenting the low tufts and thinly
scattered bushes on the edges of the meadows. This habitat differed from
that known to one of us (C.E.) in Ethiopia, where brunnescens inhabits humid
meadows with grass of a height of 0*30 to 1 m. It would seem that in this
Adamaoua locality, the grass being so short, the birds were associated rather
Hz
16000-
12000-
8000-
4000-
Hz
64000-
48000-
32000-
16000-
244
m sec
« I ■.;'.' $ I
.^.....-.....■h
27
msec.
flBtil.
27 r
Figure 1. Song in flight of Cisticola brunnescens mbangensis. A: Sonagram on standard
scale, 160 - 16000 Hz. B: The same reduced to quarter speed of A to show distribution of
energy in band 5000 - 7000 Hz. (Sonagrams made with wide band-path filter).
with the bushy formations. It may be noted that in the highlands of Bamenda
(5°5 5'N, io°io'E), in ex-British Cameroun (now part of Nigeria), where the
form C. b. lynesi Bates occurs, the species inhabits (according to the labels of
143
specimens in the British Museum, collectors Admiral H. Lynes and the Rev.
Dr. W. Serle), certainly "moorland" and "open grassy flats" but also "bare
hills with small shrubs and no new grass yet", this latter biotope resembling
that in which we found it. The males were singing, at intervals from one
another of ioo to 150 m. The song — apparently like that of the nominate
race of the Ethiopian plateaus — was performed either in flight (the most
usual) or from the top of a small bush: see sonagrams in Fig. 1. In this
habitat, Macronyx croceus and Galerida {=Heliocorys) modest a were common;
Myrmecocichla nigra and Francolinus bicalcaratus were seen ; while in the neigh-
bouring zone of high bushes the dominant species were Prima subflava,
Cisticola brachyptera, C. natalensis, Nectarinia cuprea^ Tchagra senega/a and Serinus
mo^ambicus.
Three males collected (at the finish of reproductive activity : testes 3 mm
long) have been compared with the important material in the British Museum
at Tring. It appears that this population of Adamaoua is sufficiently distinct
from the other races of the species to require a name. So we propose :
Cisticola brunnescens mbangensis, subsp. nov.
Description: Our specimens are in very worn nuptial plumage, but never-
theless are at once distinguishable from the other races in the same plumage.
They resemble most closely lynesi Bates of the highlands of Bamenda, the
only area not far from Cameroun where the species is so far known (White
1962: 683). They differ in their smaller size: see measurements below. The
general coloration is lighter. The streaking of the upperparts is less intense,
these markings being narrower and in particular more brownish, less
blackish. The crown is more rusty, not hazel. The rump contrasts with the
mantle and is a bright and dark rusty maroon, lacking the yellowish tone of
lynesi. Although very worn, the edges of the feathers of the mantle and the
wing-coverts appear more yellowish, less reddish.
Distribution: So far only known from the type locality in the highlands of
Adamaoua, Cameroun.
Type: £ adult, Adamaoua, 17th October 1972, collected by the authors,
their no. 22. In Museum National d'Histoire Naturelle, Paris (wing 50, tail
25-5, bill from skull 10 mm).
Measurements: The following are comparative measurements of the new
race and eight males in breeding dress from the highlands of Bamenda:
Wing Tail Bill from skull
C. b. lynesi
8q*0* 5I-54(52-4) 26-31(28-9) II - II- 5 (lI-2)
C. b. mbangensis
3 cTc? 49-5o(49-5) 24-26(25-1) 10-10-5(10-1)
Acknowledgements : We are grateful to Dr. D. W. Snow and I. C. J. Gal-
braith for making us welcome in the bird room at Tring and for the loan of
specimens. We are likewise thankful to C. W. Benson for translating our text
into English, and also to Mrs. J. Hall-Craggs for assistance in its finalisation.
References :
Bartholomew, J. 1956. The Times Atlas of the world. IV. London: Times Publishing Com-
pany.
White, C. M. N. 1962. A checklist of the Ethiopian Muscicapidae (Sylviinae). Part II. Occ.
Pap. Nat. Mus. S. Rhod. 26 B: 653-738.
144
The biology and a new subspecies of
Monticola sharpei
by T. Farkas
Received 29th May, 1973
INTRODUCTION
The uncertainty about the generic and specific status of rock-thrushes in
Madagascar has been discussed by Goodwin (1956). I worked in the field, in
the breeding areas of these birds, from October 1969 to January 1970, as a
result of which I was able to ascertain new details of various aspects of their
biology. These details, preceded by an examination of specimens in the
American Museum of Natural History, New York, support the suggestion
of Salomonsen (1934), Goodwin (1956) and (recently) Hall & Moreau
(1970) that the species concerned should be placed in the genus Monticola.
Indeed this had been suggested much earlier by Milne-Edwards & Gran-
didier (1879: 367), using the name Petrocincla instead of Monticola. I (Farkas
1 971) have already discussed Monticola bensoni Farkas, and I will do the same
for M. imerina (Hartlaub) in due course. The subject of the present paper is
the more complicated case of M. sharpei (Gray), which I studied in the field
as follows: — Mt. d'Ambre, 3rd November/ 3rd December; Perinet, 9th-
22nd December; Ankaratra Mts., 27th December/2nd January.
A NEW SUBSPECIES
There are several features indicating that there are three clear-cut sub-
species. For clarification I now refer only to geographical distribution and to
differences in colour-pattern in adult males. Firstly, in the highland race the
slate-blue extends from the throat to the lower part of the breast, while in
the lowland and northern (Mt. d'Ambre) races this colour does not extend
below the throat. Secondly, the margins of the remiges and wing-coverts
are rufous in the northern race; likewise in the lowland race, except that the
primaries are grey-edged; while in the highland race there are no rufous
margins except for the secondaries, which are pale rufous-edged. Thirdly, the
bright rufous rectrices are only narrowly dark-tipped in the northern race,
whereas in the other two these tips are broader, and the dark colour extends
along the outer webs of the outermost pair. Furthermore, the northern race
has a "super-rufous" phase in which the dark tips are reduced still more or
even absent, and irregular red spots may even be apparent on the dark
central pair of rectrices. Finally, in the lowland race some males have a very
pale, almost whitish patch on the chin (Plate 1), strongly reminiscent of the
patch occurring regularly in the male of M. gularis (Swinhoe) of eastern Asia.
Sharpe (1871, as Cosspyha imerina) described an adult male collected by A.
Crossley, hinting however that it might differ from the form described by
Hartlaub (i860), now known as M. imerina (Hartlaub). Thereupon Gray
(1871) designated a new species, which is now M. sharpei (Gray) (in the
References below, note the misleading title of Gray's paper, since he des-
cribed a new species o£"Cossypha" as well as o£ Capri mulgus). There is nothing
in Sharpe's description to indicate that the adult male which he had before
him does not belong to the highland race, i.e. that it has grey-edged wing-
coverts and primaries, and a slate-blue breast as well as throat. The specimen
!. is in the British Museum (Natural History), and has been examined at my
request by C. W. Benson, who confirms accordingly. It is the one listed by
145
t ,
I
si
u it*
S^i
146
Warren & Harrison (1971) as a syntype of "Cossypha sharpei G. R. Gray,
1 871" (sic), the other two syntypes doubtless being the two specimens
regarded by Sharpe (1871) as young. The size of this specimen had already
aroused the suspicion of Salomonsen (1934). He refers to it as "exceptionally
big", with a wing-length of 82, as against 75-78 mm in "eight other males
from Sianaka". Benson has also made its wing-length to be 82 mm. Thus this
specimen belongs to the blue-breasted, grey- winged, large highland race, the
correct name of which is therefore Monticola sharpei sharpei (Gray). Salo-
monsen (1934) overlooked this, and re-described the highland race as M.
imerina interioris, which is accordingly a synonym of M. s. sharpei. Conse-
quently the small lowland race is in need of a name and description : —
Monticola sharpei salomonseni, subsp. nov.
Type: £ adult, collected in May 1929, Sianaka forest, eastern Madagascar.
In American Museum of Natural History, registered number 412287.
Measurements of type: Wing 71, tail 53, culmen (from rhamphotheca) 11,
tarsus 24 mm.
Description: The adult male of salomonseni differs in colour from that of
nominate sharpei in that the slate-blue of the throat does not extend onto the
breast, and only the primaries have grey edges on their outer webs, the wing-
coverts and secondaries being rufous-edged. Salmonseni is a very small form,
indeed the smallest in the genus as a whole. For further detailed wing-
measurements, see Salomonsen (1934, in which sharpei = salomonseni,
interioris = sharpei). The tail seems disproportionately short; thus 47-59,
av. 54-8 mm, in 12 males in the A.M.N.H.
By contrast, the adult female is little more than a small version of that of
M. s. erythronotus (Lavauden), although with the strong rufous tints of this
northern (Mt. d'Ambre) race relatively subdued. Yet the most distinctive
feature is the white gular patch, reaching to the lower mandible, and edged
by a dark malar stripe. This patch is less marked in the other two races
(Plate 2). The underparts are pale brownish with a rufous tint, darkest on the
breast and its sides, where there are oblong buff spots with dark, thin edges,
diminishing downwards and replaced on the abdomen by narrow, dark
apical tips to the feathers, giving a scaly appearance. The upperparts are
plain dark olive with some red tinge, the wings similar but paler. The two
central rectrices are dark olive with very fine, dark cross-bars, the others
lighter with some rufous tinge, with a few irregularly shaped, pale rufous
spots. Such spots and the rufous tinge are less developed in the nominate
race, but more so in erythronotus, salomonseni thus being intermediate in this
respect.
Distribution: Salomonseni occupies the eastern lowland primeval forests,
down to sea-level around the Baie d'Antongil, and intergrades at higher
altitudes with the nominate race. An adult male which I measured (wing 77,
tail 57, culmen 13, tarsus 25 mm), in the museum of the Office de Recherche
Scientifique et Technique Outre-Mer (O.R.S.T.O.M.), Tananarive, seems
intermediate. It was collected in 1924 at Bejofo-Bealanana, ca. i4°3o'S.,
48°45'E. The nominate, highland race occupies the Ankaratra Mts., and high
ground on the central ridge of Madagascar to the north and south. Georges
Randrianasolo (pers. comm.) has recently collected it in the forest patches of
the Ankazobe Mts. (north-west of Tananarive), and Benson in the Massif
de l'ltremo, west of Ambositra (two females, one adult, wings 79, one
juvenile, wings 80 mm, 10/11 January). Salomonsen (1934) suggests that the
type-specimen of sharpei (that is, the syntype listed by Warren & Harrison
147
197 1 ) came from somewhere in the Sianaka forest. In fact, as Benson cor-
roborates, it is merely labelled "Madagascar". But it can be presumed that it
was collected at a higher altitude west of the Sianaka forest, perhaps in the
mountains west of Moromanga, or still further west and higher.
At present erythronotus is only known from Mt. d'Ambre, in the northern-
most tip of Madagascar. But it may intergrade to the southward with the
nominate race. According to Randrianasolo, during an O.R.S.T.O.M.
expedition in November 1966 to the Massif du Tsaratanana, led by R.
Albignac, an adult male of an unknown race of M. sharpei was collected at a
nest with two well developed young. The site was at an altitude of 2,050 m,
in light forest with many bamboos in the undergrowth, the nest being 1 • 5 m
up on a moss-covered rock. This specimen is in the collection of Col. Ph.
Milon, and still awaits determination. It could be an intermediate.
HABITAT PREFERENCE, GENERAL HABITS
In the Ankaratra Mts., from the lower edge of the primeval forests up to
2,000 m, near the summit, M. sharpei is common under the dense and high
canopy wherever there is little or no undergrowth. The floor is thickly
covered with fallen leaves and debris, in damper places with a cushion of
moss. In clearings or less dense parts of the forests there is a high and lush
shrubbery, unsuited to the species. In fact the birds prefer the more humid
parts along streams, and wooded creeks. Along such creeks I found sharpei
to be ubiquitous, although to-day most of them are cut off from the main
body of the forests, encircled by pine plantations or low, dense secondary
brush, both quite unsuited to it.
The female spends more time on the ground than does the male. There she
is perfectly camouflaged both by her cryptic colour and her quietness (Rand
1936 aptly refers to sharpei as "sedate"). Often my first sight of the female
was when she flew up only a few yards away, to alight on a low perch. If I
tried to get nearer, she flew up into the canopy. The male is also often found
on the forest floor, foraging among leaves and debris. Both sexes also take
food regularly on mossy tree-trunks, lichen-bedecked thick branches, and
even in the canopy. Their perches range from about 1 m up to the lower strata
of the canopy. If frightened, they disappear into the canopy, remaining there,
if necessary, silent and invisible for hours on end.
The race erythronotus prefers places where under a somewhat discontinuous
canopy of medium to high trees a luxuriant undergrowth of shrubs, tree-
ferns and Pandanus thrives. The damp floor is thickly carpeted with fallen
leaves, debris, moss, ferns and grass. Many strong lianas intersperse the
space under the canopy, giving variety to the typical habitat of the "hooly-
hooly", as the local foresters call these birds, in the mist-forests of Mt.
d'Ambre. Like the nominate race, erythronotus also seems attracted by streams
and creeks, although one can find it just as often in the margins as in the
depths of the forest. But it never occurs under a continuous and dense
canopy lacking in undergrowth, and never takes to places where there are
only young trees dispersed among a dense undergrowth. Subject to this, it
occurs from the lower edge of the forest, altitude ca. 600 m, to almost the
summit at 1,450 m, though mostly between 800 and 1,200 m. Assuredly this
race prefers the undergrowth more than does nominate sharpei. I often saw
males, especially at dusk, perched low by a forest path, to swoop down now
and then for an insect on the ground, or even to chase prey on the wing with
great skill. If alarmed, often with a peculiar and harsh shriek, they dived
148
with a quick loop into the undergrowth, attaining a safe distance and then
ascending into the canopy. Although I could not approach a male nearer
than some 1 5 m, sometimes I could get as near to a female as 4 m. The birds
often flew up to a high perch on a main branch from the rear, the head just
poking out, a careful watch being kept on my movements.
TERRITORY, REPRODUCTION
According to Rand (1936), the onset of breeding in most Madagascar birds
is in some way adjusted to the seasonal rainfall. On Mt. d'Ambre, while there
is ample humidity from mist throughout the year, there is also a definite
rainy season from December to March, when there are between 100 and 300
mm of rain per month. In the central highlands the picture is similar, but in
the east there is good rain in every month, except September/October, when
the monthly average can be as low as 50 mm in some places. The races
erythronotus and sharpei start breeding four to six weeks before the onset of
the rains, that is, in the second half of October or early in November. In the
east, at Perinet, I spent only the middle two weeks of December, mainly
because I soon realised that breeding there was already over. A freshly
moulted, subadult male in the A.M.N.H. (January 1923, Perinet) also sup-
ports the assumption that in salomonseni adults and young alike moult in
December/January, whereas in the other two races this starts in March. Thus,
assuming that six months are needed from time of hatching to completion of
moult out of the spotted juvenile dress, it could be that salomonseni starts to
breed in July, that is, about the driest time of the year. Possibly under the
conditions of a more evenly distributed rainfall this would be advantageous.
There is no evidence of any movement in any race of M . sharpei. In the
nominate and erythronotus territories are established in October. The largest
one (in erythronotus) I estimated to be ca. 2-5 ha, but mostly they did not
exceed 1 ha. Territorial rivalries seem to be confined to males; females, if
present, were never involved in such activities. Indication of territory being
marked by song, encroaching rivals were always received with threatening
and self-asserting postures. A rudimentary display-flight served as a nuptial
display (to the female) or as a kind of self-assertion (to another male). A male
of erythronotus ', incited by a tape play-back, performed in this manner repeat-
edly, in front of the tape-recorder. He first showed a self-asserting posture,
and then flew off to another perch, where he made a gentle bow and repeated
the self-asserting posture. The intervening flight had been straight, the wings
slowly beaten, the tail fully spread and a bright nuptial song uttered. Similar,
reduced display-flights have been reported for M. cinclorhynchus, vide Lister
(1953), and for M. gu/aris, Neifeldt (1966).
In the few cases in which nest-building was observed, only the female was
involved. Of the 18 nests found in the Ankaratra Mts., all were built on large
trees, mostly in notches on trunks, but very occasionally in forks of thick
branches, from 1-5 to 6 m above the ground. They were made entirely of
vegetable matter, the thick base and walls mainly of moss, the cup neatly
lined with fine rootlets. Typically in the highland race there is a light ring
on the rim, caused by a layer of light-coloured rootlets, those towards the
lining being darker (Plate 3). Average dimensions are: — height (including
beard-like base) 16, total width of cup 8, depth of cup 5 cm. The nest of
erythronotus is similar except for the absence of the light ring on the rim. The
fibrous lining appeared to have been obtained from lianas. However, an
important difference in site was noted. Five nests of erythronotus found were
149
Plate 3. Nest with eggs of Monticola s. sharpei; Ankaratra Mts. By author.
all skilfully hidden among Pandanus leaves, 3 to 6 m up, always in the deep
shadow of the forest canopy. A. Traka, chief forester at Les Rouselles
Station, told me that erythronotus builds only exceptionally on tree-trunks,
mostly on Pandanus. However, a sixth nest which I found on Mt. d'Ambre
had been built on the trunk of a giant tree-fern, directly over a forest path.
In four clutches of erythronotus the clutch-size was only two, only in one
was it three; the only clutch of the nominate was also two eggs. In both races
the eggs are a pale unspotted turquoise. The two eggs from the Ankaratras
measured 24 x 16-5, 23 x 17 mm. Incubation was only observed in erythro-
notus. It was carried out solely by the female; although males, feeding
incubating females, were observed at nests too. As soon as I approached a
nest, the male always sang a few quiet warning melodies, whereupon the
female slipped away from the nest. A female, two days before the hatching
of eggs, somehow failed to do so; the male then stooped down to the nest
with excited singing and chased her away. The period of incubation is not less
than 1 5 days. Both parents feed the young, which after leaving the nest are
under parental care for about one month. The nominate and northern races
are probably single-brooded ; in the lowland race the whole breeding pattern
may be different, as already suggested.
POST-EMBRYONIC DEVELOPMENT
The observations in this and the following section are based mainly on two
hand-reared young males of erythronotus which I kept from the age of 10 days
for two years in captivity (ages as given are always from date of hatching),
although supplemented by field observations. The skin of nestlings of M.
sharpei is yellow, with long, pale grey down; bill pale grey, turning blackish
150
within two weeks; legs ivory, turning blackish after six weeks. The large
head with a bright yellow gape, and long legs, are conspicuous. The sleeping
habit of nestlings is also remarkable. They lie mostly on their sides with
outstretched necks and legs, the latter "planted" against the lining of the cup
of the nest. On the nth day the eyes of my two young started to open, and
by the 1 5 th all my movements were attentively followed. On the 1 7th day
they weighed respectively 32-5, 32 g; at 500 days 41, 43 g. The first clumsy
attempts at preening started on the 12th day. They soon improved, and on
the 17th preening was extended to the wings and tail. The first head-
scratching (indirect) was observed on the 1 5 th.
In begging, the head stretched towards the feeding parent turns alter-
nately left and right from the axis of the body, following the movements of
the parent's head. The body is lifted into a sloping position while supported
on the intertarsal joints, and the wings are fluttered gently, the one nearest
to the parent usually more vigorously than the opposite one ("asymmetrical")
fluttering). "Symmetrical" fluttering also occurs, but vanishes soon after
leaving the nest. The first vocal component is a quick succession of weak
"peeping" tones, changing gradually after the 24th day into a humming
"sreee", rather similar to the begging calls in other Monticola spp. This call,
used in begging or for any other purpose, disappears when the young can
feed independently.
The droppings are large, pear-shaped and enveloped in a tough, gelatinous
substance. After leaving the nest they become smaller, and the gelatinous
envelope disappears. My hand-reared young spent all their time either free
in my room, or in a cage when I was on the move. But during a difficult stage
of the expedition in January 1970, for reasons of their security, at night they
were hung in linen bags over my bed. They accommodated themselves to this
with such success that in the mornings they would only defaecate after I
opened the bags. But the droppings were again as they had been in the nest-
ing stage.
On the 19th day the young left the nest. They moved around on the ground
with quick hops as do adults. (Young in general seem to spend the first few
weeks mostly on the ground. Those which I encountered in the Ankaratras
in December, when disturbed, quickly flew into the canopy, descending
again to the floor when it seemed safe to do so.) On the 24th day the first
attempt was made to pick up ants from the ground; by that time they were
able to fly some distance with ease, and even make turns in mid-air. The first
attempt to catch an insect in flight was made on the 30th day. By the 40th
they were dismembering grasshoppers and earthworms, and within a few
days had become self-supporting, although begging did not completely
disappear until the fourth month.
On leaving the nest, the male can be identified by the vivid red tail, and the
brighter lemon-yellow soles and orange gape (in the female the gape and
soles are by contrast pale white, this contrast being a widespread feature in
the genus as a whole, at all ages). By the 40th day, when the wings and tail
had attained their full length, sexual dimorphism becomes further marked by
the start of the growth of the axillaries, orange in the male, white tinged
yellow in the female. Moult out of the juvenile dress started about the 120th
day, on the underparts and nape, reaching its climax in the fifth month and
ending early in the sixth. The subadult male has the full colour of the adult.
The latter has apparently only one moult a year, and there is no marked
151
seasonal difference in plumage in M. sharpei. However, subadults can be
distinguished from adults quite easily. This is because in the juvenile dress
the greater wing-coverts have buffy apical markings and the secondaries pale
buffy edges. These feathers, together with the primaries, axillaries and
rectrices, are not changed in the post-juvenile moult.
I was obliged to cut-back the primaries of my hand-reared young of M.
sharpei (and also those of young Copsychus albospecularis pica obtained at an
early stage of the expedition), in order to prevent their escape. Also, owing
to crowded cages and constant travelling, some tail-feathers were broken.
After my return to my home in South Africa, I left these feathers alone,
hoping that I could gradually remove them. But to my surprise, when the
moult came, all the damaged primaries and rectrices were replaced in a
natural way, while the undamaged ones were retained. A few other hand-
reared M . imerina and C. a. inexpectatus which I collected at a later stage of the
expedition, and which escaped cutting or damage, moulted normally; i.e.,
they retained all their remiges and rectrices. Thus it seems that functional
inadequacy caused by damage as described above can induce premature
moulting in the young.
DEVELOPMENT OF VOICE AND BEHAVIOUR PATTERNS
Adults of M. sharpei warn with a quiet "hjutt-tock-tock". Such notes can
also be heard sometimes separately. The "hjutt" was first heard from the
young on the 1 8th day, when they were frightened by a sudden movement on
my part. The "tock" was first heard on the 21st day, and accompanied by a
bow. A distress-call, the shriek already referred to, was also first heard on the
21st day. The displeasure-call, a staccato "kerrr-errr", was only first heard on
the 117th day; on the same day the social-call was also first heard — a gentle
"weed", repeated at longer intervals, serving as mutual contact between the
sexes, and often heard during the field observations.
First attempts at subsong were made on the 17th day, while sleeping in the
nest. The throats of the young pulsated, and a few weak inarticulate notes
were heard. On the 20th day they sang while awake. Thereafter the daily
amount of time spent in this exercise gradually increased, but even by the
40th day there had been no increase in volume. The rasping, guttural
double tones, so typical of the rehearsed songs of all Monticola spp. (Greenewalt
1968), were first heard on the 59th day, increasing graduallv so that about the
1 30th day they had become a general feature of the song of these two young.
During the seventh month subsong was completely superseded by the re-
hearsed song, which in adults of the genus is the prevailing form of song in
the off-season, though it can also be heard in the breeding season, restricted
to the hottest time of day. This rehearsed song in M. sharpei is always of a
very low volume, although rich in the aforementioned guttural notes and in
imitations. Development of motij 'or primary song also started at an early stage.
On the 64th day one of the young burst into a few, very loud tones (a car had
started near us); on the 90th day they repeatedly sang at dusk a few semi-loud
motifs, and on the 120th day, again at dusk, they rendered a few short
melodies. By the 280th day their melodies were in no way inferior to those of
adult males; the volume of their primary song remained, however, generally
low with a marked daily peak at dusk, until about the 350th day, when the
volume of song increased considerably within a few days, heralding the
onset of reproductive activity. There is a marked difference in the primary
songs of at least M. s. erythronotus and M. s. sharpei, which, if this were
152
considered alone, could suffice to regard them as specifically distinct. The
repertoire oi erythronotus consists of a dozen or so highly stereotyped melodies
while that of the nominate race is wider and characterized by a measure of
variation between individuals. Singing at dusk is typical of both races, but I
never heard singing at night.
As regards the function of primary song in M. sharpei, the above features
distinguish the territory song of the races erythronotus and sharpei. Like so many
other features, the song of salomonseni awaits investigation. The signal song,
being only a shortened form of territory song, serves warning purposes in
both races. The limited song-repertoire of females was only heard in this
context; thus as soon as I approached a fledged young, it was mainly the
female which uttered a few weak signal-motifs, coupled with the staccato
note of displeasure and threatening postures.
The courtship song can be directed by the male either at a female or another
encroaching male. In either case it is co-ordinated with courtship/self-
asserting postures, and consists entirely of tonal material peculiar to the
species but with an unmistakeable similarity to the courtship songs of other
Monticola spp. In the young it was first heard on the 196th day.
Both the nominate race and erythronotus can imitate. But, while erythronotus
does only in the rehearsed song, sharpei imitates in both the rehearsed and
territory songs. They imitate only the calls, never the songs, of other birds.
The first imitation was heard on the 131st day. It was of the call of Malaco-
notus ^eylonus, a common species around my home in the Cape. The reper-
toire was further enlarged on the 196th day by imitation oiParus afer; after
the juvenile moult, calls of Nectar ini a famosa, Prinia maculosa and Spreo hicolor
were added. On the 220th day imitation was heard of Hartlaubius auratus and
Tchitra mutata, and in the following weeks of Coracopsis vasa and Nesiilas
typica, which had last been heard some five months previously while still in
Madagascar.
In addition to begging as described above, other fixed action patterns
became apparent, in sequence as follows: — The twitching of the folded wings is
only a gentle movement of the remiges including the primaries. It was first
observed in sleeping nestlings only 16 days old. On leaving the nest, a slow
tilting of the folded tail (up to ca. 30 degrees) was noted. While wing-twitching
can be repeated several times in quick succession, tail-tilting is always a
single movement. On the 9 1 st day tivitching of the folded tail was noted — a
very quick spreading-and-folding of the rectrices. It is not very conspicuous
because only the two outermost pairs of feathers are used in this scissor-like
movement. It can be repeated, but only after a long interval. The tail can also
be completely spread (fanned) and remain in this position for a while. Finally,
the fanned tail can also be twitched by a very swift folding-and-spreading of the
two outermost pairs of feathers only.
Two variations of head-movement were also noted in M . sharpei. On one,
the head is pointed motionless towards the opponent, and the neck somewhat
intracted. In the other, the head is tossed rhythmically to the left-straight
ahead-to the right, thence back to straight ahead. In both variations the head
feathers are kept firmly depressed.
The sole intention movement is performed immediately on leaving the nest.
The clumsily hopping young, attaining some elevated point, assumes an
upright stance and makes a single, lazy bow synchronized with wing-
twitching and tail-tilting. In great excitement only, this intention move-
ment can be repeated twice or thrice in quick succession. The interesting
153
point is that this movement soon fades away with age; adults seldom so
perform.
Interspecific aggressiveness was first noted on the 38th day, i.e. playful
chasing of a fledged Copsjchus albospecularis. On the 43 rd day the first action
of intraspecific aggressiveness followed. On the 5 9th with all feathers de-
pressed and the body erect, the young showed the first greeting ceremony; the
head pointed slightly upwards and the bill slightly open, a rapid subsong was
uttered. From this juvenile posture the courtship display was gradually
evolved.
On the 96th day the first threatening posture was noted; the feathers on the
mantle, back and lower breast somewhat bristled, the head pointed stiffly
towards the opponent, the wings and tail twitched synchronously at irregular
intervals and a few warning calls uttered (this posture was observed in both
sexes). Soon thereafter followed a stage of development when the young
often showed ambivalent threatening and greeting postures while singing
primitive courtship songs. From these elements first on the 280th day the
self-asserting posture ("imponieren") and then on the 300th day the courtship
display evolved. In the self-asserting posture all the feathers are depressed,
and the tail widely spread and either titled up (if the male stays on the floor),
or pressed down (if on a branch). In either position the tail is also twitched
repeatedly. The head is tossed rhythmically, and a few sharp whistles uttered
(cf. M. rupestris and M. angolensis, in Farkas 1962, 1968). If self-asserting
is performed on the floor, the male runs around with clumsy steps ; if on a
branch or other perch, he makes several turns, showing himself off alternately
from the front and the back to his opponents (males), which keep their dis-
tance. The courtship display is almost the same except that it is accompanied
by a courtship song typically mingled with "tock-tock's" and mimicry. If the
male performs on the floor, he again runs around the female.
AC KNOWLEDGEMENTS
The present paper is the third in a series of studies on Madagascar thrushes
(for the first two, see Farkas 1971, 1972), a project which has been fully and
most generously supported by a grant from the Frank M. Chapman Memorial
Fund of the American Museum of Natural History. During the visits to Mt.
d'Ambre, the Ankara tra Mountains and Perinet I enjoyed the assistance and
hospitality of the Department des Eaux et Forets of the Government of the
Malagasy Republic and of the O.R.S.T.O.M., Tananarive. Father Otto
Appert, C. W. Benson (particularly), Professor J. Berlioz and G. S. Keith
kindly provided me with valuable information while preparing the manu-
script. I am most grateful for all their support.
SUMMARY
The status of the polytypic Monticola sharpei is reviewed, and a diagnosis
given of its lowland race. In support, details of morphology, distribution and
habitat preference are compared for the three subspecies so far known. The
possibility of the existence of a fourth one is also mentioned. While in plu-
mage, breeding season, habitat preference, primary song, nest and choice of
site thereof, and timing of moult, there are certain divergences between the
subspecies, as regards eggs and clutch-size, incubation, care of offspring,
fixed action patterns, and calls and rehearsed songs, there seem to be no
marked differences.
154
References :
Farkas, T. 1962. Zur biologie und ethologie der siidafrikanischen arten der gattung
Monticola (Boie). II. Teil: Monticola rupestris L. Die Vogelwelt 83: 161-173.
— 1968. Zur biologie und ethologie der Angola-Waldmerle, Monticola angolensis Sousa.
Rev. Zool. Bot. Afr. 77: 162-189.
— 1 97 1. Monticola bensoni, a new species from south-western Madagascar. Ostrich, suppl.
9: 83-90.
— 1972. Copsychus albo specular is winterbottomi, a new subspecies from the south-east of
Madagascar. Ostrich 43: 228-230.
Goodwin, D. 1956. Note on the genus Pseudocossyphus Sharpe. Bull. Brit. Orn. CI. 76:
143-144.
Gray, G. R. 1871. On a new species oiCaprimulgus. Ann. Mag. Nat. Hist. (4)8: 428-429.
Greenewalt, C. H. 1968. Bird song: acoustics and physiology. Washington, D. C. : Smiths. Inst.
Hall, B. P. & Moreau, R. E. 1970. An atlas of speciation in African passerine birds. London:
Brit. Mus. (Nat. Hist.)
Hartlaub, G. i860. Systematische uebersicht der vogel Madagascars. II. Journ. Ornith. 8:
81-112.
Lister, M. D. 1953. Secondary song of some Indian birds, fourn. Bombay Nat. Hist. Soc.
52:1-40.
Milne-Edwards, A. & Grandidier, A. 1879. Hi stoire physique, naturelle et politique de Mada-
gascar. 12. Paris: Imprimerie Nationale.
Neifeldt, I. 1966. The wood "Rock-thrushes". Biol. Ptici Trudi Zool. Inst. Nauk. Leningrad
39: 120-184.
Rand, A. L. 1936. The distribution and habits of Madagascar birds. Bull. Amer. Mus. Nat.
Hist. 72(5): 143-499-
Ripley, S. D. 1964. Family Muscicapidae : Subfamily Turdinae, Thrushes. In Check-list of
birds of the world. 10: 13-272. Cambridge, Mass.: Mus. Comp. Zool.
Salomonsen, F. 1934. On a hitherto unknown fauna of montane birds in central Mada-
gascar. Nov. Zool. 39: 207-215.
Sharpe, R. B. 1871. Contributions to the ornithology of Madagascar. II. Proc. Zool. Soc.
Lond: 313-320.
Warren, R. L. M. & Harrison, C. J. O. 1971. Type-specimens of birds in the British Museum
{Natural History). 2. London: Brit. Mus. (Nat. Hist.)
The natal pterylosis of the Swallow-tanager
by Charles T. Collins
Received 3rd July, 1973
The natal pterylosis of passerine birds has received some attention in North
America (Wetherbee 1957, 1958) and more recently in South Africa (Markus
1 971). However, such information is nearly lacking for the vast assemblage
of Neotropical birds, other than unquantifled field observations included in
various life history studies and brief quantified observations made on two
species of tanagers (Collins 1963). Data are particularly lacking for species,
as the Swallow-tanager Tersina viridis, which make up some of the smaller
and distinctive taxa inhabiting this region. Until recently this distinctive
fruit eating Neotropical species has been placed in the monotypic family
Tersinidae. More recently (Storer 1970) it, and the tanagers of the closely
related family Thraupidae, have been relegated to subfamilial status within
the more inclusive family Emberizidae. However, Sibley (1970, 1973) con-
sidered Tersina to be only a distinctive genus in the tribe Thraupini and
probably most closely related to Thraupis and Tangara.
The information presented here on the natal pterylosis of Tersina viridis
was obtained principally from eight preserved specimens of young birds. All
were collected by me in the vicinity of the Estacion Biologica de Rancho
Grande, Est. Aragua, Venezuela during the spring of 1972, and are referable
155
to T. v. occidentatis. Of these eight, five were very small nestlings (Stage A,
see Wetherbee 1957: 356) and three were late stage embryos removed from
eggs which expectedly would have hatched within a day or two. The ad-
ditional data from three nestlings w T hich were examined in the field before
being returned to the nest are incomplete in that the small neossoptiles of
the ventral tract, secondaries and rectrices were overlooked and thus not
recorded. The data from all other tracts have been included since they are in
close agreement with the more detailed counts made on the preserved
specimens.
The total number of neossoptiles varied from a low of 77 to a high of 107
per bird. However, four of the eight specimens had from 101 to 103 neos-
soptiles and thus a typical number of 102 can be assigned to the species based
on this and the number most frequently observed for each individual tract.
As discussed later, two tracts were exceptionally variable and for one of these
the numerical average was used since there was no single typical number for
the tract. The total number of neossoptiles is given in Table 1, and the
distribution of these neossoptiles in Tersina is given in Table 2.
Table 1. Total number of neossoptiles
Species Number Total number of
of specimens neossoptiles
Tanagra (E upborn a) violacea 1 32*
Thraupis palm arum 2 184-224*
Piranga olivacea 2 227**
Tersina viridis 8 102 (77-107)
* from Collins (1963)
** from Wetherbee (1958)
Table 2. Counts of neossoptiles of Tersina viridis
Tract Number of neossoptiles
Minimum
Typical
Maximum
Coronal
3
4
6
Occipital
4
4
5
Humeral
6
8
10
Spinal (upper)
4
5
7
Spinal (lower)**
2
8*
12
Femoral
4
7
8
Ventral
6(5*)
9
Secondary
1
1
Secondary Coverts
4
6
7
Caudal
6
6
6
* numerical average to nearest whole number
** unpaired tract along midline; all others paired
The only previous information on the natal pterylosis of Tersina is the
statement by Schaefer (1953: 439) that they have "sparse natal down". The
down of all of the specimens examined in this study appeared to be uniformly
whitish or greyish-white, whereas Schaefer reported it to be "of the same
yellowish-pink color as the rest of the body, giving the skin a peculiar light
golden hue". This discrepancy is not easily resolved as both studies were
conducted in the immediate vicinity of the Estacion Biologica de Rancho
Grande.
The number of neossoptiles for each tract usually varied within rather
narrow limits and was constant or nearly so in some. Exceptions to this were
the lower midline portion of the spinal tract and the ventral tract. In the
former case the great variation (2 to 12 neossoptiles) made the choice of a
typical number impossible and the average number was utilized. In the case
of the ventral tract four specimens had four, five or seven neossoptiles but
others had less (in one case none) and one had nine. A compromise number
156
would seem to be six: a value not actually observed in any specimen. The
average number of neossoptiles in the ventral tract of all specimens was five.
If this average value is used as the typical number for the tract the typical total
count for the species would be reduced to ioo, a value slightly less than the
most frequently observed total. Thus the choice of six as the typical number
for this tract seems more appropriate. The neossoptiles from most tracts
were of medium length as compared to those of other species. However, the
neossoptiles of the ventral tract, secondaries and rectrices were very short
(1-2 mm) and, as mentioned above, overlooked in the course of field ob-
servations of one brood of nestlings. The total number of neossoptiles is
noticeably less than for two of the three species of tanagers for which data
are available (Table i). Both of the species of tanagers having higher total
counts of neossoptiles build normal open nests while Tersina nests in bur-
rows (Schaefer 1953) and Tanagra (Euphonia) violacea builds a compact domed
nest. Further information is needed on a wider array of tanagers and their
allies (Collins unpublished) before much weight can be given to this seeming
correlation between "cavity" nesting and a reduction in number and dis-
tribution of neossoptiles.
Acknowledgements : The specimens of Tersina utilized in this study were
collected as part of a continuing programme of study of Neotropical birds
supported by research grants from The Frank M. Chapman Memorial Fund
of the American Museum of Natural History, New York. I am indebted to
Dr. Gonzalo Medina Padilla for permission to stay at Rancho Grande and
for the permit to collect material for this study. Manuscript preparation was
aided by a Summer Faculty Fellowship from the California State University,
Long Beach Foundation.
References :
Collins, C. T. 1963. The natal pterylosis of tanagers. Bird-Banding 34: 36-38.
Markus, M. B. 1970. A preliminary survey of the occurrence of neossoptiles in South
African passeriform birds, with special reference to natal pterylosis. M.S. Thesis, Univ.
of Pretoria (available from Univ. Microfilms, Ann Arbor, Mich., U.S.A.; Masters
Abstr. 9[ 3 ], 1971).
Schaefer, E. 1953. Contribution to the life history of the Swallow Tanager. Auk 70: 403-
460.
Sibley, C. G. 1970. A comparative study of the egg-white proteins of passerine birds. Bull.
Peabody Mus. Nat. Hist. 32: 1-131.
— 1973. The relationship of the Swallow-tanager Tersina viridis. Bull. Brit. Orn. CI. 93:
75-79-
Storer, R. W. 1970. Subfamilies Thraupinae and Tersininae. In Check-list of birds of the world.
13. Cambridge, Mass.: Mus. Comp. Zool.
Wetherbee, D. K. 1957. Natal plumages and downy pteryloses of passerine birds of North
America. Bull. Amer. Mus. Nat. Hist. 113: 339-436.
— 1958. New descriptions of natal pterylosis of various bird-species. Bird-Banding 29:
232-236.
A small collection of birds from Muara Island,
north Borneo
by Kenneth IV. Prescott
Received 1 6th July \ 19J)
Reading Max C. Thompson's account (1966) of birds which he had collected
on north Borneo caused me to refer to my notes concerning a small group of
specimens which I had collected from Muara Island, north Borneo in June
and July 1945. At first reading, it appeared that he had collected most of the
157
species on the eastern coast of north Borneo but not all that I had found off
the west coast. Recent comparison of my Muara birds with specimens at the
American Museum of Natural History and with those at the United States
National Museum disclosed a new subspecies of iora (Prescott 1970b), a
kingfisher race new to Borneo (Prescott 1970a) and an important record of
the Stork-billed Kingfisher in Borneo (Prescott 1972). It seems advisable,
even after such a long post-collecting period, to cause a brief account of this
small collection, the only known (to me) from Muara, to be added to the
literature.
Muara is a small island in Brunei Bay located off the north-west coast of
Borneo just north of the Sarawak boundary, at approximately 5°N longitude
and 1 1 5 °E latitude. Collecting was carried out with the co-operation of the
United States Navy as the writer was then serving as Executive Officer of the
U.S.S. Jamestown, AGP-3. The specimens, made up as skins, are in the col-
lection of the Museum of Zoology, University of Michigan (UMMZ), Ann
Arbor, Michigan. The following species account is based on field notes of
three brief collecting trips made during morning hours.
Treron curvirostra nasica, Thick-billed Green Pigeon: Specimen, 1: juv.,
sex(?), 17th July 1945, UMMZ 114,000. Several were observed on the ground
and in thick bushy undergrowth, two or three feet high, alongside a small
fresh- water stream about 50 feet from the ocean beach. They were shy and
when frightened flew a short distance upstream to the protection of similar
bushy growth. Thompson (1966: 395) collected T. c. curvirostra and Smythies
(1968: 232) gives T. c. curvirostra as resident throughout Borneo although
questioning if this subspecies is recognizable as the form on Banggi and
Balambangan Islands.
Halycon chloris co/Iaris, White-collared Kingfisher: Specimens, 2: male and
female, 17th July 1945, UMMZ 113,926 and 113,927. The only two seen
were collected near the ocean shore ; the female from a tall deciduous tree and
the male from low second growth. These are the first recorded specimens
from Borneo (Prescott 1970). Thompson (1966: 401) collected H. c. chlorop-
tera and this is the race reported by Smythies (1968: 308-310) as resident
throughout Borneo in coastal areas and estuaries, and a familiar garden bird
in Brunei. It is of interest that Smythies (Joe. cit) found evidence of seasonal
movement for H. c. chloroptera, i.e. being present during the breeding season
(January-August) in the west coast Kimanis Bay area just above the range of
the Stork-billed Kingfisher but absent thereafter. Although I did not see H.
c. ch/oroptera, my specimens were taken from the same general locality in
which I took a Stork-billed Kingfisher.
Pelargopsis capensis innominata, Stork-billed Kingfisher : Specimen, 1 : sex
(?), nth June 1945, UMMZ 114,018. This specimen has already been fully
commented on by Prescott (1972: 160).
Lalage nigra scbisticeps, Pied Triller: Specimens, 2: male and female (?),
17th and 19th July, UMMZ 113,994 and 113,995. Both specimens were taken
from low lying bushy, second growth. The species was seen but not taken
by Thompson (1966: 416). Smythies (1968: 380) describes L. n. nigra as the
race in Borneo and the north Bornean Islands and Maratuas. Chasen & Kloss
(1930: 60) collected eight specimens of L. n. nigra on an island lying off the
north-eastern coast of Borneo.
Aegithina tiphia trudiae, Common Iora: Specimens 4: two males and two
females, 17th and 19th July 1945, UMMZ 113,996 through 133,999. This
group represents a new race (Prescott 1970b: 39-40). Thompson (1966: 420)
collected individuals of A. v. viridissima in south Borneo and Sarawak and
158
A. t. aequanimis in north Borneo and Palawan. However, Smythies (1968:
387-388) gives A. t. viridis for south Borneo and Sarawak, A. t. aequanimis
for north Borneo with the boundary between it and viridis coinciding roughly
with the "Sarawak-British North Borneo" boundary.
Anthreptes malacensis bornensis, Brown-throated Sunbird: Specimens, 6:
four females and two males, nth June, 17th and 19th July 1945, UMMZ
114,002 to 114,006 also 114,019. All were collected from flowering bushes
in low second growth at the edge of a cleared area. Individuals returned to
the same area several minutes after a shot was fired and were in company
with Aethopyga s. siparaja. Thompson (1966: 427) also collected this species.
Smythies (1968: 492-493) gives A. m. bornensis for north Borneo including
the north Bornean islands and A. m. malacensis for south Borneo and the
Karimatas.
Aethopyga s. siparaja, Yellow-backed Sunbird : Specimen, 1 : male, 1 7th
July 1945, UMMZ 114,007. In company with the last species, several were
seen in the flowering bushes at the edge of a clearing. They were extremely
shy and difficult to approach. Smythies (1968 : 498-499) describes the species
as a common resident in the lowlands of the main island of Borneo, Banggi
Island, the Maratuas and Karimatas. However, Thompson (1966: 428)
described three siparaja individuals taken at Tawau which he noted as having
darker wings than others of the race.
Dicaeum trigono stigma, Orange-bellied Flowerpecker: Specimen, 1: juv.,
female (?), 17th July 1945, UMMZ 114,001. Collected from the flowering
bushes at the wood's edge with the two sunbirds (above). Identification of
this juvenile to subspecies is uncertain. Thompson (1966: 431) collected one
specimen of V. t. dayakanum. Smythies (1968: 487) gives D. t. trigonostigma
from the Karimatas and D. t. dayakanum from the main island of Borneo.
Acknowledgements : I am grateful to Max C. Thompson and Robert W.
Storer who kindly read an early draft of this note and made critical suggest-
tions; moreover, the latter kindly loaned the UMMZ Muara specimens. I
shall always be indebted to the late Josselyn Van Tyne who encouraged me
to make the South Pacific collections.
References :
Chasen, F. N. & Boden Kloss, C. 1930. On a collection of birds from the lowlands and
islands of north Borneo. Bull. Raffles. Mus. 4: 1-112.
Prescott, Kenneth W. 1970a. A kingfisher new to Borneo. Auk 87: 586.
— 1970b. A new subspecies of the Common Iora from north Borneo. Bull. Brit. Orn. CI.
90: 39-40.
— 1972. Pelargopsis capensis innominata in north Borneo. Bull. Brit. Orn. CI. 92: 159-160.
Smythies, B. E. 1968. The birds of Borneo. 2nd. ed. Edinburgh & London: Oliver & Boyd.
Thompson, Max C. 1966. Birds from north Borneo. Univ. Kansas Publ. 17(8).
Coracina papuensis in Indonesia
by C. M. N. White
Received 20th August, iyj)
Coracina papuensis is a cuckoo shrike which ranges from Australia and New
Guinea to the Solomon Islands and the eastern Indonesian Archipelago.
In the last area Peters (i960: 1 79-1 81) lists two forms: melanolora Gray in the
northern Moluccas, and timorlaoensis Meyer in the Tenimber Islands.
Melanolora hardly merits recognition as it is like the nominate form of west
and north New Guinea but averages larger. Mees (1972: 81-82) has most
recently pointed out that it is an extremely poor subspecies. In the case of
159
timorlaoensis the distribution given is incomplete and the validity of the
subspecies questionable.
Meyer described timorlaoensis 'in 1884 from two specimens from the Tenim-
ber Islands, both females. No other collector has obtained it there, including
Kiihn who made a considerable collection there at the end of 1900 and in
early 1901. Finsch (1901: 250) listed a female from Kisar collected by
Schadler on 4th December 1897, and still in the Leiden Museum. Finsch
identified this as the north Australian form hypoleuca, of which he considered
timorlaoensis a synonym. He also reported a female from Great Kei collected
on 23rd May 1865 by Hoedt, which he likewise identified as hypoleuca. Kiihn
made large collections in the South- West Islands, including Kisar, but the
only occasion when the species was found was in June 1906, when two fe-
males were collected on Sermatta. Hartert (191 1: 166) identified these as
timorlaoensis with a query and a remark that it was doubtful if the form could
be distinguished from hypoleuca. Kiihn also made large collections in the Kei
Islands where he did not obtain the species. Van Bemmel (1948) overlooked
the Great Kei record, and Peters (i960) has overlooked it and also the records
from Kisar and Sermatta.
Thus there is little to support the existence of a distinct subspecies,
timorlaoensis, in the Tenimber Islands. Rather there are a few scattered
records, all of females, from Tenimber, Great Kei, Sermatta and Kisar which
seem identical with north Australian hypoleuca. Indeed they are perhaps only
occasional nomadic migrants, as the dates in May and June, the Australian
winter, may indicate. The December record in that case is peculiar. In the
Northern Territory in Australia hypoleuca is known to breed from November
to January.
I am grateful to Dr. G. F. Mees for information about the specimens in
the Leiden Museum, and for discussing the question dealt with in this note.
References :
Bemmel, A. C. V. van. 1948. A faunal list of the birds of the Moluccan islands. Treubia 19:
323-402.
Finsch, O. 1901. Systematiche uebersicht der vogel der Sudwest Inseln. Notes Leyden Mus.
22: 225-309.
Hartert, E. 191 1. On the birds of Sermatta. Nov. Zool. 18: 161-167.
Mees, G. F. 1972. Die vogel der Insel Gebe. Zool. Meded. 46: 69-89.
Peters, J. L. i960. In Check-list of birds of the world. 9. Cambridge, Mass.: Mus. Comp.
Zool.
Vachycoccyx audeberti: some addenda
by C. W. Benson & AL P. Stuart Irwin
Received 1 8 th fitly, 1973
We (1972, Arnoldia, Rhodesia 5(33): 24 pp.) detailed what is known about the
distribution, biology and systematics of this cuckoo. A few points worth
mention have come to light subsequently.
On p. 3 we drew attention to its sparseness, both in Africa and in Mada-
gascar. Further emphasis to this is lent by the fact that in the original des-
cription of nominate audeberti (Schlegel 1879, Notes Leyden Mus. 1 : 99) it was
reported that the example collected by Audebert was the only one met with
in the course of two years in north-eastern Madagascar.
On p. 8 we were un able to accept a record of parasitisation of the Violet-
backed Starling Cinnyricinclus leucogaster (see originally Neuby-Varty 1946,
160
Ostrich 17: 345). Neuby-Varty gave the measurements of the egg claimed to
be of this cuckoo as 25 x 18 mm. The Rev. Dr. William Serle (pers. comm.)
has pointed out that the colour, markings and si%e are correct for C. leuco-
gaster, and that it is most unlikely to be an egg of Pachycoccyx. Furthermore,
as he writes, the broken egg of this cuckoo reported by Fischer & Reichenow
(1879, J ourn - Ornith. 27: 341) (see our p. 6) was only slightly longer but no
wider than an egg of Euplectes flammiceps (=hordeaceus). McLachlan & Liver-
sidge (1970, Roberts birds of South Africa, 3rd ed., Cape Town: John Voelcker
Bird Book Fund: 564), for instance, give an average measurement for the
egg of E. hordeaceus as 1 8 • 5 x 13-8 mm only. Indeed so far the only proven
host is the Red-billed Helmet Shrike Prionops retail.
We regret that on p. 14 no acknowledgement was made to Dr. Kenneth C.
Parkes, Curator of Birds at the Carnegie Museum, Pittsburgh. He most
kindly lent Benson the specimen in that museum (p. 19). See p. 20, Dr. J.
Steinbacher reports that there is no specimen in the Natur-Museum und
Forschungsinstitut Senckenberg, Frankfurt.
We did not use a male in the Transvaal Museum from "Chambere, G. E.
Afr., 24.9.03, J. V. O. Marais", since we could not trace this locality (p. 19).
However, Mrs. M. K. Rowan and Dr. A. C. Kemp have drawn our attention
to a note by Roberts (1935, Ostrich 6: 48), in which he discusses the specimen
in that museum from the Olifants River, 16th July 1934, and another one
therein from "near Bagamoyo, Tanganyika Territory". Evidently Roberts
had knowledge of the history of this specimen, not recorded on the label,
and it may reasonably be presumed to be one and the same as the Chambere
specimen. We have still been unable to trace Chambere on any map, but it
can be accepted as being near Bagamoyo, in coastal Tanzania, and whence we
recorded (p. 18) a specimen in the Museum of Comparative Zoology,
Harvard. The Chambere specimen was examined by Irwin. Although Roberts
found the difference between it and the Olifants River specimen "quite
pronounced", and attributed the former to P. a. validus (Reichenow), the
latter to P. a. canescens Vincent, we attribute both to validus, of which
canescens is a synonym, and founded on a specimen in very fresh dress (see
our p. 12). The Chambere specimen has wing 217, tail 182, culmen from base
29 mm.
Dr. Kemp reports that the Transvaal Museum has recently received a
female specimen collected by O. P. M. Prozesky at Croc Ranch, Phalaborwa
area, eastern Transvaal, at 23°5 5'S., 30°5 5'E., 1,500 ft. a.s.l., 10th September
1972; wing 242, tail 202, culmen from base 27- 3 mm. Also, Kemp himself saw
an individual on the farm Charleston, eastern Transvaal, at 24^ 3'S., 3 1°3 5'E.,
1,000 ft. a.s.l., 1 2th July 1973. It was moving steadily from tree to tree in dry
leafless mixed woodland, but was not seen to feed.
Some remarks on the Lammergeyer Gypaetus barbatus
in East Africa
by E. Sch'u^
Received 20th July, 1973
In his interesting bird observations on the Shira Plateau and Kibo, Kiliman-
jaro, King (1973, Bull. Brit. Orn. CI. 93: 68) gives only one record of the
Lammergeyer, an individual of which he saw on 6th November 1968. Guest
& Leedal (1954, Tang. Notes & Recs. 36: 47), who King quotes, and who
161
spent two months on the mountain, never saw it. However, there are other
records from Kilimanjaro, to which attention should be drawn. Fuggles-
Couchman & Elliott (1946, Ibis 88: 330) saw a pair west of the Shira ridge
on 24th October. Schiiz (1958, Altes und Neues iiber das Vorkommen des
Bartgeiers {Gypaetus barbatus) in Ostafrika. Journ. Ornith. 99: 394-398) gives
at least five records since 1932 from various localities on Kilimanjaro, apart
from several by the missionary, the late Dr. Reusch. G. H. H. Brown (1963,
Journ. E. Afr. Nat. Hist. Soc. 24: 72) gives another record — an immature
bird which accompanied his safari on the march between the Bismarck and
Peters Huts. So far the only material ("in the hand") evidence is from a
feather, a secondary, found by me (Schuz 1958, op. cit.: 396) on 5th Septem-
ber 1957 at Peters Hut (3,800 m), now in the bird exhibition rooms at Schloss
Rosenstein (Staatl. Mus. fur Naturkunde in Stuttgart). The Game Depart-
ment report of an immature Lammergeyer caught and photographed by
Wilhelm von Furstenberg (foot-note in Moreau 1944, Tang. Notes <& Recs.
18 : 48) is obscure, since Baron Furstenberg wrote to me in 1958 that he only
had one field observation and had never possessed a specimen of this species
or photographed it.
As mentioned (Schuz 1958, op. cit.: 397), records of the Lammergeyer in
East Africa are of interest as snowing a connection between the centres of
distribution in Ethiopia in the north and the Drakensberg Mts., in Natal, in
the south. The most southerly outlying northern (East African) record is
from Mbeya, southern Tanzania (Moreau pers. comm. to Schiiz 1958, op.
cit. : 397), and the most northerly outlying southern (South African) one from
Maruma (20°2 5'S., 33 E.) on the Mozambique/Rhodesia border. The
Maruma record is based on a specimen (Swynnerton 1908, Ibis [9] 2: 425).
Both these records were repeated by Benson & Irwin (1963, Ardea 51 : 222).
With regard to the Maruma one, Irwin (pers. comm., 4th August 1973)
informed Benson that he knows of no further record from either Mozambique
or Rhodesia, and suggested that it probably represented a migrant from
further south. Benson has also written to me as follows: — "Despite the fact
that ornithologically it has been well explored and documented — Belcher
(1925, Ibis [i2]i : 797-814) and Vincent (1934, Ibis [i3]4: 126-160) are only
random examples — I know of no record of the Lammergeyer from Mt.
Mlanje, southern Malawi. Mlanje rises to ca. 3,000 m, the uppermost 1,000 m,
above the montane grasslands, being bare and boulder-strewn. There is an
excellent account of this mountain by Chapman & White (1970, The evergreen
forests of Malawi. Oxford, Commonwealth Forestry Institute: 1 62-1 71)".
To bring my synopsis of East African records (Schiiz 1958, op. cit.) up to
date, the following references are relevant: — Hale (1958, Journ. E. Afr. Nat.
Hist. Soc. 23 : 16); L. H. & G. H. H. Brown (1963, Journ. E. Afr. Nat. Hist.
Soc. 24: 72); Paul Geraghty (1969, Animals 12: 312); Dewhurst (1971, E.
Afr. Nat. Hist. Soc. Bull. : 133); and Gerhart (1971, E. Afr. Nat. Hist. Soc.
Bull.: 173). Brown & Brown (1963, op. cit.) claim that their records from
Mt. Elgon and Muruanisagar, 40 miles west of Lodwar, thus both slightly
west of 3 5°E., appear to be a considerable westward extension of the known
range in East Africa. However, records from the Ngorongoro Crater, in
northernT anzania (Schiiz 1958, op. cit. : 397; Huxley & Nicholson 1963, Ibis
105 : 106) are from almost as far west, at ca. 3 5°3o'E.
Brief reference may be made to what little is known about the breeding
status of the Lammergeyer in East Africa, partly based on information from
L. H. Brown (pers. comm. to Benson, 6th September 1973). North (1948,
Ibis 90: 1 3 8- 1 41) has given some information from Kenya, but there still
162
seems to be no known breeding site in Tanzania. North records an egg taken
at a site within 50 miles of Nanyuki in May, and according to Brown the site
was in the Loldaiga Hills, near Nanyuki. North also mentions a site within
50 miles of Naivasha, but could not give a date for breeding. According to
Brown a pair started to breed in the Njoroma Gorge (Hell's Gate), near
Naivasha, in 1964, and has done so every year since, except 1971. This site
has been visited by up to 2,000 tourists annually. Information therefrom is
the source of what is written in Brown & Amadon (1968, Eagles ; hawks and
falcons of the world. London, Country Life Books: 313), that egg-laying takes
place in Kenya in January; likewise that from the Nanyuki site in May.
North also briefly mentions a site on Mt. Kenya, and Coe (1967, Monogr. Biol.
17. Den Haag: 108) implies that the Lammergeyer breeds on that mountain.
There must surely be breeding sites in Tanzania and other ones in Kenya.
Further information is required, but the most urgent necessity is protection.
I am much indebted to C. W. Benson and Dr. L. H. Brown for their
assistance in the finalisation of this paper.
Organochlorine insecticide residues and food remains in
a Bald Ibis Geronticus eremita chick from Birecik, Turkey
by J. L. F. Par slow
Received 24th August, 197)
The Bald Ibis Geronticus eremita is a rare, declining species whose only known
breeding colonies are in Morocco and on a cliff in the town of Birecik, which
stands beside the upper Euphrates in south-east Turkey (see Smith 1970).
The only other member of the genus is the almost equally rare G. calms (also
known as the Bald Ibis and regarded by some authorities as conspecific with
eremita) which is restricted to South Africa where there are about 1,000 pairs
in 70 known breeding colonies (Siegfried 1971). The numbers of eremita are
still decreasing. The total known population of wild birds in 1972 was only
about 150 pairs, of which about 60 pairs bred; 26 of these nested at Birecik,
in which area there were as many as 530 pairs in 1953, but only 130 pairs in
1962 and 45-48 pairs in 1967 (Kumerloeve 1962, 1965, 1967; Porter 1973;
U. Hirsch in litt.). The species is listed in the I. U.C.N. Red Data Book of
endangered animals, and studies of its breeding ecology in Turkey have
recently been initiated under the auspices of the World Wildlife Fund as
part of a programme of conservation measures. These studies are being
carried out by U. Hirsch wha (in litt.) has traced the decline to two main
factors : first, the uncontrolled use of DDT ,dieldrin and other insecticides in
the Birecik area by the Turkish Ministry of Health during 1958-60, which
resulted in more than 100 Bald Ibises being found dead in 1959-60 in the
colonies and in the fields in which the birds feed, and second, the increased
disturbance of the colonies due to house building in the immediate vicinity.
Kumerloeve speculated that increased cultivation along the banks of the
Euphrates (and the consequent loss of feeding areas), as well as poisoning
from synthetic insecticides, had caused the decline. In order to determine
whether the synthetic organochlorine insecticides might still be having an
effect on the birds, a chick and an egg collected in 1972 were sent to Monks
Wood Experimental Station for analysis. This paper discusses the results and,
because so little detailed information exists on the species' diet, also reports
on the food remains found in the chick's stomach.
163
CHEMICAL ANALYSIS OF AN EGG AND CHICK
The egg was collected from the colony by U. Hirsch, C. D. W. Savage and
others on 4th May 1972. The contents showed no signs of embryonic
development. The chick was seen to fall from a nest (during a quarrel with
one of its siblings) on 15 th May 1972 and was picked up dead beneath the
colony. It was about 10-15 days old. The egg contents and liver of the chick
were analysed at Monks Wood for organochlorine insecticide residues using
gas-liquid chromatography. A sample of the food remains from the stomach
of the chick was also analysed for organochlorine residues but with negative
results. In addition, the egg contents were analysed by atomic absorption
spectrophotometry for certain potentially toxic metals. Determinations for
cadmium (Cd) and lead (Pb) were made but were negative (levels of detection,
ca. o- 2 ppm). Positive results are given in Table 1.
TABLE 1
Organochlorine and toxic metal concentrations (in parts per
Ibis chick and egg. (ND = none detected, <o.o5 ppm; * =
million wet weight)
not analysed.)
in a Bald
pp'-DDE
dieldrin
mercury
(Hg)
copper
(Cu)
zinc
(Zn)
Egg i-8
Chick • 4
0-3
ND
o- 12
*
1-34
*
15-8
*
In avian tissue, DDT almost invariably metabolises to DDE, and aldrin to
dieldrin. The finding of DDE in the egg and chick and dieldrin in the egg
indicates that the parent birds must have been exposed, presumably through
their prey, to DDT and aldrin/dieldrin at some stage during their life, if not
in Turkey then on migration or in their winter quarters in north-east Africa.
The residues found, however, are not especially high and are unlikely at these
levels to be having serious sublethal effects — that is unless the Bald Ibis
happens to be very much more sensitive to these materials than any other
bird species yet tested. Certainly the concentrations recorded are equivalent
to the normal 'background' contamination levels now present in the eggs and
tissues of many bird species in Britain.
The concentration of toxic metals detected in the egg are probably also
normal. Hg is present in small quantities in virtually all British bird tissues
and eggs, and in seabird eggs for example are normally 4 to 50 times higher
than the levels in the Bald Ibis egg without causing any apparent adverse
effects. Symptoms of Hg poisoning in gallinaceous species do not occur
until liver residues reach at least 1 5 ppm (Borg et al. 1969), while reduced egg
hatchability in the same group does not occur until tgg residues reach about
o- 5 ppm (Fimreite et al. 1970). Cu and Zn, in trace quantities, are of course
essential for all forms of life and the concentrations reported here are in no
no way atypical for an avian egg.
FOOD REMAINS IN THE CHICK
Both Geronticus species appear to have similarly varied diets, though no
detailed studies have been published. Various insects and other terrestrial
invertebrates, molluscs, lizards, frogs, fish and dead small birds and mammals
are all mentioned in the literature as prey items (e.g. Siegfried 1966, Smith
1970, Porter 1973), while Pala (1971) found that the stomach contents of a
Birecik adult caught on 5 th June 1969 comprised almost entirely locusts
164
and bush-crickets (70 % Calliptamus italicus, 2 5 % Dociostaurus maroccanus,
5 % Tettigoniidae) plus a large, hairy spider. On dissection at Monks Wood,
the stomach of the chick from Birecik was found to contain the fragments of
a wide range of mainly ground-living insects. A representative sample was
used for the organochlorine analysis (see above) and the remainder (about
two-thirds of the total) were sent to the British Museum (Natural History)
for identification. At least 15 different insect species were represented (48
individuals) belonging to seven families; five crustacean claws were also
present. A full list of species with minimum numbers of individuals of each
is given in Table 2.
TABLE 2
Food remains in the gizzard of a Bald Ibis chick from Birecik
Species
Min. no. of
individuals
Gryllopalpidae (mole-crickets)
Gryllotalpa gryllotalpa (L.) adults :
4
nymphs :
11
Formicinae (ants)
Messor sp.
1
Camponotus sp.
11
Tenebrionidae (nocturnal ground beetles)
Amnodeis }confluens Mill.
1
Adesmia procera Mill.
1
Pimelia bajula Klug
5
indet. sp. (small)
1
Carabidae (ground beetles)
Zabrus rotundicollis (Men.)
2
Scarites sp.
1
Buprestidae (jewel beetles)
Julodis onopordi (F.)
2
Scaraboidae (chafer and dung beetles)
Copris ?hispanus L.
3
}Scarabaeus sp.
2
indet. (2 spp.)
2
Curculionidae (weevils)
indet. sp.
1
Crustacea
indet. no. of claws :
5
ACKNOWLEDGMENTS
Colleagues at Monks Wood Experimental Station — M. C. French, A. A.
Bell and Dr. R. C. Welch — were responsible for the chemical analyses and
other technical assistance, and Peter Hammond at the British Museum
(Natural History) identified the prey remains from the chick; U. Hirsch and
C. D. W. Savage made valuable comments on an earlier draft of this paper.
I am grateful to them all.
References :
Borg, K., Wanntorp, H., Erne, K. & Hanko, E. 1969. Alkyl mercury poisoning in ter-
restrial Swedish wildlife. Viltrepy 6: 301-379.
Fimreite, N., Fyfe, R. W. & Keith, J. A. 1970. Mercury contamination of Canadian prairie
seed-eaters and their avian predators. Can. Fid Nat. 84: 269-276.
Kumerloeve, H. 1962. Zur geschichte der Waldrapp-kolonie in Birecik am oberen Euphrat.
Journ. Ornith. 103: 389-398.
— 1965. Zur situation der Waldrapp-kolonie Geronticus eremita in Birecik am Euphrat.
Vogelwelt 86: 42-48.
— 1967. Nouvelles donnees sur la situation de la colonie d'Ibis chevelus Geronticus
eremita (L.) 1758 a Birecik sur l'Euphrate (Turquie). Alauda 35 : 194-202.
165
Pala, Sukru. 1971. Faydali kuslarimizdan ( Geronticus — Comatibis — erewita L.) — ozellikleri,
yasayisi ve beslenmesi. Tabiat ve Insan 5 : 14-22.
Porter, R. 1973. The disappearing ibis. Birds 4: 227-228.
Siegfried, W. R. 1966. The Bald Ibis. Bokmakierie 18: 54-57.
— 1971- The status of the Bald Ibis of southern Africa. Biol. Conserv. 3: 88-91.
Smith, K. D. 1970. The Waldrapp Geronticus eremita (L.). Bull. Brit. Orn. CI. 90: 18-24.
Nesting of Verreaux's Eagle Aquila verreauxi in Zambia
by J. F. R. Colebrook- Robjent <& T. 0. Osborne
Received 29th August, 1973
The breeding of Aquila verreauxi is well known from Tanzania, Malawi and
Rhodesia, but has not so far been recorded from Zambia. In Benson et al.
(1971) there are sight records of this eagle from rocky hills in the Eastern and
Northern Provinces. It has recently been seen by R. J. Dowsett and Cole-
brook-Robjent in the gorges below the Victoria Falls in the Southern
Province: see the cyclostyled Bull. Zambian Orn. Soc, June 1972, 4(1): 25.
Nyanje Hill is particularly mentioned by Benson et al., and it is from this
area that we now record the first known nest of this eagle in Zambia. On
nth June 1972 Colebrook-Robjent and R. Stjernstedt discovered a nest in a
wind-hole cave about halfway up a sheer 70 m granite rock face on Mbewa
Hill, west of Nyanje Village. One of the pair was flushed from the nest,
which was judged to contain eggs. One of us (J. C.-R.) attempted, but did
not reach the nest. On 19th May 1973 J. C.-R. and T. O. O. visited the same
nest-site, and from the opposite side of a ravine we saw the eagle on the nest.
Later T. O. O. reached the nest, which contained two eggs. The large stick
nest almost covered the "cave" floor, measuring approximately 2- 5 wide by
1 • 5 m high. The cup was lined with green but dry Brachystegia leaves and
measured 45 cm across. The two bluish white eggs were fairly evenly, but
lightly spotted, and measured 77-1 X 60 • 2, 75 • 4 x 58-4 mm. They weighed
respectively 150 and 1 3 5 g. One was collected and proved to be almost fresh.
The egg-white was sent to Prof. Charles Sibley of Yale University for
analysis.
Brown & Amadon (1968) give the average measurements of 30 eggs as
76-9 X 58-6 mm. The peak egg-laying month in Rhodesia is May, and the
only such record from Malawi is also for May (see Benson et al. 1964, as
augmented for Rhodesia by Vernon 1965).
A glance at any detailed map of the area will suggest that there are other
pairs of Verreaux's Eagles inhabiting the scattered rocky hills along the
southern boundary of the Eastern Province of Zambia. Similar habitat
extends far into neighbouring Mocambique.
References :
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. S. 1971. The birds of Zambia.
London: Collins.
Benson, C. W., Brooke, R. K. & Vernon, C. J. 1964. Bird breeding data for the Rhodesias
and Nyasaland. Occ. Papers Nat. Mus. S. Rhod. 27B: 30-105.
Brown, L. & Amadon, D. 1968. Eagles, hawks and falcons of the world. London: Country Life
Books.
Vernon, C. J. 1965. The 1964 Black Eagle survey in the Matopos, Rhodesia. Arnoldia
{Rhodesia) 2(6): 9 pp.
166
Observations on Bradypterus carpalis and
Brady pterus graueri
by Ruth Trimble Chapin
Received ist September, 1973
(1) Bradypterus carpalis Chapin
The original series of Bradypterus carpalis (four males; one female) was
secured in the Uele District of the then Belgian Congo from 191 1 to 191 3
and was described in 1916. The species was next recorded by van Someren
(19 1 9). Apparently without knowledge of the description of carpalis, van
Someren named his birds jokanae. Delacour (1943: 34) and Chapin (1953:
435) consi&crz&yokanae not even subspecifically distinct from carpalis.
For many years these were the only known specimens of this species, and
it came to be regarded as one of the rare birds of Africa. However, Schout-
eden (1966) recorded a specimen taken in the valley of the Akanyaru River,
east of Butare (formerly Astrida) Rwanda; and more recently Britton &
Harper (1969) reported collecting seven specimens of carpalis near Lake
Kanyaboli, western Kenya.
West Kivu volcanoes
MumbaB J
Nyawarongo"*/ KiW
LwiroT^ B Rugege Forest
Akanyaru R«J ^ ' /""'
TANZANIA
28
Figure 1 : Localities of specimens of Bradypterus graueri (■) and B. carpalis (#). Open
circle represents field observations of the latter.
No additional specimens from the Congo (now called Republique du
Zaire) were obtained until the species was rediscovered in a marsh near
Lwiro, Kivu District. The sedge warblers were always of great interest to
167
my husband, and at every likely marsh we watched and listened for Braqyp-
terus of any kind. At Tshibati, 6,400 ft., Kivu District, where we lived for five
years, we found only B. baboecala centralis. But at Lwiro, more than a thousand
feet lower in altitude, we heard on numerous occasions a distinctive chirping
song which my husband attributed to B. carpalis. Its secretive habits always
made it a difficult bird to observe, but it is rather easily recognized by its
large size, dark brown upperparts, the black markings on the breast, and the
white shoulder patch which is evident in flight. In a papyrus swamp in
Queen Elizabeth Park, Uganda, along the Kazinga Channel, we watched
and listened to carpalis in April 1956. In south western Uganda, just west
of Kabale, again in papyrus at 6,200 ft., we heard the song repeatedly in
December 1956 and again in February 1957. It was also heard many times in
a similar area 45 miles southwest of Mbarara, at 4,800 ft., but the bird could
not be seen.
Eventually through the efforts of Boniface Mulimbwa, our Congolese
assistant, eight specimens (five males ; three females) were collected from the
marsh near Lwiro, at 5,200 ft. With Boniface we had verified the presence of
carpalis in this marsh and directed his attention to its song. Although carpalis
is usually associated with papyrus swamps, the habitat of the original speci-
mens and those collected in Uganda and Kenya, the present series was taken
from elephant grass and broad-leaved sedges. In the same marsh we also
collected B. baboecala centralis, and its presence there provided an oppor-
tunity to compare the calls of the two species, which are well described by
Chapin (1953: 43 3> 436).
When comparing the Lwiro specimens of carpalis with those of the type
series, one can observe a difference in their appearance. The underparts of
the Lwiro birds are whiter, and the black markings on the chest seem heavier
and more extensive. The tone of brown on back and tail is deeper in the more
recently collected specimens. The measurements of the Lwiro birds exceed
those of the birds from the Uele. While the two populations seem to differ,
the size difference is small and that in colour perhaps in part at least is the
result of foxing. The opportunity to compare our Kivu birds with recent
material was provided by Mr. A. D. Forbes-Watson of the National Museum
of Kenya, who kindly sent two specimens from the Britton/Harper series
taken at Lake Kanyaboli, Kenya. The specimens from Kenya and Lwiro are
comparable in colour; they differ only in size. Measurements in millimetres
follow: —
Culmen Hindtoe
Male Wing Tail from base Tarsus & claw
Type series
(Uele Distr.) 67-72 69-75 18-19 27-28 18-21-5
Type oiyokanae
(now in AMNH) 70 73 20 28 21
L. Kanyaboli, Kenya 70-72 73-76 18-18-5
Lwiro, Kivu Distr. 73-76 76-78-5 18-5-19-5 29-30 19-21
Female
Type series
(Uele Distr.) 68 67 18 26 18
L. Kanyaboli, Kenya 66-69 66-74 17-18-5
Lwiro, Kivu District 71-73 74-76 18-19 27-5-29 19-21
We have weights for only two of the birds taken at Lwiro: male, testes
enlarged, 12th Dec. 1957, 27 g; female, ovary somewhat enlarged, 12th Dec.
168
I 95 7> 2 5 g- The measurements of the Kenya specimens are from Britton &
Harper (1969).
Sedge warblers like Bradypterus seem always to be in worn plumage or
moulting, so that small differences in wing and tail measurements, such as
these, do not justify recognition by a subspecific name. This material, how-
ever, extends the range oi carpalis to the Kivu and establishes that its habitat
is not exclusively papyrus.
(2) Bradypterus graueri Neumann
The relationship of B. graueri, carpalis, and grandis has been somewhat
uncertain. White (i960: 430, footnote) maintained that all three are con-
specific, with graueri as the oldest name. Britton & Harper (1969) and Britton
(1970) have accepted his opinion by writing of carpalis as a race of graueri.
Hall & Moreau (1962, 1970), however, following Chapin (1953) and Rand
et al. (1959), "regard all three as distinct, though related species, forming a
superspecies". This opinion seems to be supported by the acquisition of more
material in recent years.
The American Museum of Natural History, N.Y., now has three addi-
tional specimens of graueri from near the type-locality, collected by J. P.
Chapin just north of Alimbongo, Kivu District, at 7,200 ft.; and an adult
male from the Rugege Forest, Rwanda, taken at 7,550 ft., 67 km west of
Butare (formerly Astrida). Dr. A. Prigogine has secured a dozen specimens
from the mountains west of Lake Kivu at altitudes from 7,086 to 7,546 ft. for
the Musee Royal de l'Afrique Centrale (Schouteden 1969 and Prigogine
in lift.). In the same museum is another specimen from the Rugege Forest,
Rwanda (Schouteden 1969). Friedmann & Williams (1968) reported 12
examples from the Impenetrable Forest, Uganda, secured in mist nets at
6,750 ft.
Although not as yet found together, graueri and carpalis occur in Rwanda
at localities that are separated by less than 50 miles, but at different altitudes.
The Bwindi Swamp of the Impenetrable Forest, where graueri was secured,
is not much farther from Kabale, Mbarara, and the Kazinga Channel, where
we have heard carpalis on several occasions.
In its relatively longer, more graduated tail of twelve feathers, and the lack
of a white shoulder-patch, adult graueri is unlike carpalis in appearance.
Britton (1970) has described the immature of carpalis as having the under-
pays "dusky brown, being especially dark on the flanks, sides of breast,
vent and legs, and entirely lacking the adult's white centre to breast and
belly". This contrasts markedly with the description of an immature graueri,
quoted from Chapin's manuscript notes: "13 Aug. 1959. Prigogine sent also
one Bradypterus graueri, male juv. (skull still soft to touch) taken at Nyawa-
rongo [Kivu District] on 24 Feb. 1959. It looked to me at first like B.
baboecala, because of the narrower spots on chest; but it has wing 62 mm.,
tail 70 mm., culmen to base 13 mm. It is thus too large for baboecala. The
light superciliary line goes back behind the eye as in graueri, and it has no pale
yellowish tinge on throat as in juvenile baboecala [our specimen from Lwiro,
5,500 ft., 25 June 1957]. But there is the faintest suggestion of a pale yellow
tint on lower breast. I doubt that I have ever seen a young graueri before."
An immature male in our Kivu collection from near Alimbongo, 5 th Oct.
1955, does retain some pale yellowish feathers on the lower abdomen, which
is mainly white; the flank feathers are cinnamon brown.
Dr. Prigogine has kindly arranged for me to re-examine the "subadult
male" referred to above. It has no evidence of spots on the throat, and the
169
chest-markings are narrower and less pronounced than those of the young
male in our collection. His specimen may be wrongly sexed, because the adult
male and female (in breeding condition) in our series do exhibit such a
difference; in fact the throat of the female is almost pure white. This con-
forms to Chapin's earlier observation (1953: 434): "In the male the spots
extend to the throat, but there they are much smaller in the female''' (italics mine).
Friedmann & Williams (1968: 29) find no such sexual difference in their
specimens, but have misinterpreted Chapin's statement.
I am grateful to Dean Amadon of the American Museum of Natural
History for reading this paper and providing helpful suggestions. Other
colleagues in the Department of Ornithology have also given advice and
assistance; but I am indebted most of all to James P. Chapin, who taught me
to know and love the birds of Central Africa.
References :
Britton, P. L. & Harper, J. F. 1969. Some new distributional records for Kenya. Bull.
Brit. Orn. CI. 89: 162-165.
Britton, P. L. 1970. The immature plumage of two African warblers. Bull. Brit. Orn. CI.
90: 26-28.
Chapin, J. P. 1916. Four new birds from the Belgian Congo. Bull. Amer. Mus. Nat. Hist.
35: 23-29.
— 1953- The birds of the Belgian Congo. Part 3. Bull. Amer. Mus. Nat. Hist. 75a.
Delacour, J. 1943. The bush- warblers of the genera Cettia and Bradypterus, with notes on
allied genera and species. 2nd part. Ibis 85 : 27-40.
Friedmann, H. & Williams, J. G. 1968. Notable records of rare or little-known birds from
western Uganda. Rev. Zool. Bot. Afr. 77: 11-36.
Hall, B. P. & Moreau, R. E. 1962. A study of the rare birds of Africa. Bull. Brit. Mus.
{Nat. Hist.) Zool. 8: 316-378.
— 1970. An atlas of speciation in African passerine birds. London: Brit. Mus. (Nat. Hist.)
Rand, A. L., Friedmann, H. & Traylor, M. A. 1959. Birds from Gabon and Moyen Congo.
Fieldiana: Zool. 41: 219-41 1.
Schouteden, H. 1966. La faune ornithologique du Rwanda. Mus. Roy. Afr. Centr. Doc.
Zool. 10.
— 1969. La faune ornithologique du Kivu. II. Passereaux. Mus. Roy. Afr. Centr. Doc.
Zool. 15.
Someren, V. G. L. van. 191 9. Exhibition and description of new birds from Africa. Bull.
Brit. Orn. CI. 40: 19-28.
White, C. M. N. i960. A check list of the Ethiopian Muscicapidae (Sylviinae) Part 1. Occ.
Pap. Nat. Mus. S. Rhod. 24B: 399-430.
A Discography of Bird Sound from the Oriental
Zoogeographical Region
by Jeffery Boswall
Received 14th September, 1973
This paper is the fifth to be published of a number of projected papers, each
covering a major geographical region of the world. For the Palearctic
Region see Boswall (1964, 1966, 1969a, 1969b, 1970, 1971) and Sellar
(1973); for Australasia see Boswall (1965); and for the Ethiopian Region,
Boswall & North (1967); and for the Antarctic, Boswall & Prytherch (1969).
A Neotropical discography is under way (Boswall & Freeman in prep.) and
it is hoped soon to complete global coverage with a Nearctic discography.
170
In the Oriental Region the first bird recording was probably that of a
hornbill {Toe k us sp.) made by the Coolidge-Carpenter expedition (Carpenter
1940). Since then, a number of workers have recorded sound production by
birds in a number of countries within this region. For example, Burton
(1969, 1 971) includes in his catalogue a few recordings by Ivan Polunin
from Malaya, Singapore and Borneo; also from Borneo Lord Medway's
recordings of Low's Swiftlet Collocalia maxima. More recently, the BBC
Sound Archives (J. F. Burton pers. comm.) brought in some recordings
made by Philip Wayre on the Bhutan/Assam border, and some from Malaya
by T. C. White. Also, BBC sound recordist John Davies made a number of
recordings in Borneo in June 1972, and Lyndon Bird, also of the BBC,
recorded a number of birds in Indonesia and East Malaysia (Borneo) in
February, March and April 1973. It is hoped that some of these will also
eventually be included in the BBC's Sound Archives.
The commonness and portability of tape recorders must mean that many
ornithologists have recorded sound production by birds in the Oriental
Region over the last ten or fifteen years. It is beyond the scope of this paper
to try to trace private collections. A few examples will suffice. L. P. Ferdi-
nand (pers. comm.), a member of the Danish "Nooma Dan" expedition,
taped about twenty species in the South Phillipines. Tom Harrison (pers.
comm.), operating from the Sarawak Museum in Sarawak, made a number of
recordings — especially hornbills, owls and bulbuls in the 1950's. M. A.
Bradshah of Madras in India started a collection in 1966. Lord Fermoy made
a number of recordings during the expedition to Pakistan led by Mountfort
(1969). D. V. Manning presented recordings of some Malayan bird calls to a
meeting of the British Ornithologists' Club on 23rd January 1968 (not 1967
as printed in Bull. Brit. Orn. CI. 88: 21).
The discography: The series of papers of which this is one is primarily con-
cerned with commercially published gramophone records. For the Orient
there are only three : —
1. Teeuwen, Gus. 1970. Jungle Memories. One 12-inch 33-3 r.p.m. disc,
no. LP 0808. Singapore: Guson (obtainable from: P.M. Films, 26 Penn
Meadow, Stoke Poges, Bucks, England).
Twenty-two bird species can be heard on this record, along with two
mammals, an amphibian and a chorus of cicadas. Of particular beauty is the
song of the Common Shama, and the incredibly pure tones arranged in a
melody by the Plain Babbler. It is interesting to hear a recording from the
wild of that common cage bird, the Hill Myna. Lengthy, well-informed
introductions are spoken by Arthur Gick.
Side 1 : Malaysian Eared Nightjar Eurostopodus temminckii ; White-handed Gibbon Hylobates
lar; Cicada chorus Cicadiidae; Common Shama Copsychus malabaricus ; Long-tailed Parakeet
Psittacula longicauda; Hill Myna or Tiong Gr acuta religiosa; Plain Babbler Malacopteron affine;
Large Racket-tailed Drongo Dicrurus paradiseus: Little Barbet Megalaima australis; Large-
footed Wren Babbler Napothera macrodactyla ; Black Hornbill Anthrafoceros malayanus;
White-throated Bulbul Criniger phaeocephalus.
Side 2: Black Crested Magpie Platysmurus leucopterus ; Frog sp. Rana erythraea; Serpent
Eagle Spilornis cheela; Red-rumped Tree Babbler Stachyris maculata; White-bellied Black
Woodpecker Dryocopus javensis ; Red-headed Tailor Bird Orthotomus sepium; Black-necked
Tree Babbler Stachyris nigricollis ; Scrub Bulbul Criniger bres ; Indian Cuckoo Cuculus microp-
terus; Black Siamang Hylobates syndactylus ; Chestnut-capped Laughing Thrush Garrulax
mitratus; Red-bearded Bee-eater Nyctiornis amicta; Muller's Barbet Megalaima oorti; Large
Hawk Cuckoo Cuculus sparverioides.
171
2. Teeuwen, Gus. 1970. Jungle Fascination. One 12-inch 33-3 r.p.m. disc,
no. LP 0809. Singapore: Guson (obtainable as for disc 1.)
The twenty-two recordings include sixteen birds, four mammals, a reptile
and a chorus of amphibians. The Common Shama and the Black Siamang are
repeated from the disc reviewed above.
Side 1: Black Siamang Hylobates syndactylus ; Mountain Imperial Pigeon Ducula badia;
Yellow-crowned Bulbul Pycnonotus ^eylanicus ; Abbott's Jungle Babbler Trichastoma abbotti;
Grey-bellied Squirrel Callosciurus caniceps ; Bronzed Drongo Dicrurus aenus ; Pygmy Owlet
Glaucidium brodiei; Rhinoceros Hornbill Buceros rhinoceros; Golden Tree Babbler Stachyris
chrysaea; Large Scimitar Babbler Pomatorhinus hypoleucos; a gecko Gecko stentor.
Side 2 : Common Shama Copsychus malabaricus ; Banded Leaf Monkey Presbytis melalophos ;
Little Cuckoo Dove Macropygia ruficeps; Ashy-naped Tailorbird Orthotomus cucullatus ; Red-
headed Laughing Thrush Garrulax erythrocephalus ; Frog spp. Phrynella pollicaris and others;
Long-billed Partridge Rhi^pthera longirostris ; Malay Peacock Pheasant Polyplectron malacenis ;
Fire-tufted Barbet Psilopogon pyrolophus ; Long-tailed Nightjar Caprimulgus macrurus; Tiger
Panthera tigris.
3. Teeuwen, Gus. 1971. Tarn an Negara {Malaysia's National Park). One
7-inch 45 r.p.m. disc, no. SP 0810, incorporating 20 page booklet. Singapore:
Guson (obtainable as for disc 1).
Twelve birds sing from this record. There is no human voice. Nor,
incidentally, is there any indication of the speed at which the record should
be played; however, by comparing the recording of Abbott's Jungle Babbler
on this record with the same species on the second disc above (side 1), it is
clear that this disc should be played at 45 r.p.m. This most helpfully devised
little publication includes maps, botanical photographs, a bibliography and
hints to park visitors !
Side 1 : Magpie Robin Copsychus saularis ; Abbott's Jungle Babbler Trichastoma abbotti;
Red-rumped Trogon Harpactes duvauceli; Cicada Cicadiidae; Paradise Flycatcher Terpsi-
phone paradisi; Great Argus Pheasant Argusianus argus; Yellow-crowned Bulbul Pycnonotus
^eylanicus.
Side 2: Changeable Hawk-Eagle Spi^aetus cirrhatus ; Malaysian Black-headed Oriole
Oriolus xanthonotus ; Slender-billed Crow Corpus enca; Common Shama Copsychus malabaricus ;
Black Crested Magpie Platysmurus leucopterus , Bushy-crested Hornbill Anorrhinus galeritus.
R eferences :
Boswall, Jeffery. 1964. A discography of Palearctic bird-sound recordings. Brit. Birds 57,
special supplement : 1-63.
— 1965. A catalogue of tape and gramophone records of Australasian Region bird-sound.
Emu 65 : 65-74.
— 1966. New Palearctic bird-sound recordings in 1964-65. Brit. Birds 59: 27-37.
— 1969a. New Palearctic bird-sound recordings during 1966-67. Brit. Birds 62(2) : 49-65.
— 1969b. New Palearctic bird-sound recordings during 1968. Brit. Birds. 62(7): 271-281.
— 1970- New Palearctic bird-sound recordings during 1969. Brit. Birds 63: 324-332.
— 1971- New Palearctic bird-sound recordings during 1970. Brit. Birds 64: 431-434.
Boswall, Jeffery & Freeman, W. P. in preparation. A discography of bird-sound from the
Neotropical Zoogeographical Region.
Boswall, Jeffery & North, Myles, E. W. 1967. A discography of bird-sound from the
Ethiopian Zoogeographical Region. Ibis 109: 521-533.
Boswall, Jeffery & Prytherch, Robin J. 1969. A discography of bird-sound from the
Antarctic. Polar Record 14(92): 603-612.
Burton, John. 1969. Catalogue of Natural History Recordings, BBC.
— 1 97 1. First Supplement to the 1969 Catalogue of Natural History Recordings, BBC.
Carpenter, C. R. 1940. A field study in Siam of the behaviour and social relations of the
Gibbon Hylobates lar. Comparative Psychology Monographs 16(5): 1-2 12.
Mountfort, Guy. 1969. The Vanishing Jungle. London: Collins.
Sellar, Patrick (1973). New Palearctic bird-sound recordings in 1971-72. Brit. Birds 66:
303-510.
172
Rufous phase Senegal Coucals in southwest Nigeria:
an illustration of Gloger's Rule
by J. H. Elgood
Received nth September, 197$
There is no doubt that the percentage of dark rufous Senegal Coucals
Centropus senegalensis, formerly regarded as a distinct species C. epomidis,
diminishes with distance inland from the Nigerian coast, a diminution
paralleled by a decrease in the average rainfall.
Although there were several earlier suggestions that the dark "Rufous-
bellied Coucal, Centropus epomidis Bonaparte 1850" was only a melanic form
of the Senegal Coucal, C. senegalensis (L.) 1 766, this view was only finally
accepted when Elgood (1955) produced evidence of interbreeding of the two
forms at Ibadan. Thus Hall & Moreau (1962), quoting Elgood, listed
epomidis in an appendix of rejected species, while White (1965) stated cate-
gorically that epomidis is a rufous form of senegalensis.
My observations on the dark form at Ibadan showed that 16% of all
observed coucals were of the epomidis type. During a recent visit to Lagos
University, in early 1972, it was found that 28 out of 65 coucals (43 %) were
rufous birds. The most northerly known records of the epomidis form are
from Oyo and Iwo, both about 50 km north of Ibadan. Further north still,
at Ilorin, 150 km north of Ibadan, no melanic individual was ever observed
during a number of extensive visits to the area, where the normal form of
senegalensis is abundant.
Unfortunately no strictly comparable rainfall data are available, but the
following is a summary of the facts : —
Station
Distance
from coast
Rainfall
belt
" epomidis
coucals
Lagos University
Ibadan
10 km
120 km
1 50-200 mm
100-150 mm
43%
16%
Oyo and Iwo
Ilorin
170 km
270 km
100-150 mm
50-100 mm
present
absent
If one postulates a direct relationship between percentage occurrence of
dark birds and rainfall, as indicated by distance from the coast, the data from
Lagos and Ibadan suggest epomidis will occur only to a distance of about
200 km from the coast, a hypothesis fully supported by the findings at Oyo,
Iwo and Ilorin.
However sparse the data, it seems clear that the percentage of rufous
coucals in southwest Nigeria diminishes rapidly from the humid coastal belt
as rainfall decreases progressively inland. Although Gloger's Rule, as
most often formulated, states that average pigmentation is heavier in
populations from more humid areas, lighter in those from more arid ones
it is only an extension of this concept to consider the proportion of dark
forms instead of average individual darkness, as a manifestation thereof.
References :
Elgood, J. H. 1955. On the status of Centropus epomidis. Ibis 97: 586-587.
Hall, B. P. & Moreau, R. E. 1962. A study of the rare birds of Africa. Bull. Brit. Mus.
(Nat. Hist.) Zool. 8(7).
White, C. M. N. 1965. A revised check-list of African non-passerine birds. Lusaka: Govern-
ment Printer.
173
The eggs of Carpococcyx renauldi
by Michael Walters
Receeved 1 2th October, 1973
Recently four eggs of Renauld's Ground Cuckoo Carpococcyx renauldi, of
southern and eastern Siam and Indochina, were presented to the British
Museum (Natural History), Tring, by L. W. Hall, of "Birdland", Bourton-
on-the-Water. The eggs, which had not been incubated, were laid by a
captive female in the summer of 1973. They are oval and tapered slightly
at one end, resembling the egg of a domestic fowl in shape, except one egg
which is a pure oval. They are white, with a rough-textured surface and are
not glossy. They measure respectively 61 • 3 x 43-7; 6o- 8 x 44-3; 62-8 x
45 • 3 ; and 62-8 X 45*5 mm.
The only reference to the eggs of this species in the literature appears to
be in Meise (1964). A single tgg in the Rothschild collection at Tring
measuring 44-4 X 34 *o mm is referred to. This egg cannot at present be
traced, but its identity would now seem to be in doubt.
Reference:
Meise, W. (ed.) 1964. Schonwetter: Handbuch der oologie. Berlin.- Akademie Verlag. 10: 593.
The Common Snipe Gallinago gallinago in Zambia
by E. L. Button
Received 27th October, 1973
Benson, Brooke, Dowsett & Irwin (1970, Amoldia, Rhodesia 4(40): 13) quote
a statement by White (1945, Ibis 87: 466) that I collected Gallinago gallinago
at Ndola, Zambia*. But in the absence of a preserved specimen in the Academy
of Natural Sciences, Philadelphia, to which I sent many specimens, or
apparently anywhere else from Zambia, they were unable to accept the oc-
currence of this species in the territory as being certain. This decision was
duly reflected by the same authors (1971, The Birds of Zambia. London:
Collins), in which the species was placed in brackets.
Major Melvin A. Traylor (pers. comm., 15th Feb. 1973) has confirmed to
me that there are six specimens of G. gallinago in the Field Museum of Natural
History, Chicago, collected by myself in the Itawa Swamp, Ndola, viz. : z<$<$ y
1 6th/ 1 8th Jan. 1944; $, 24th Feb. 1944; <$, 2nd Mar. 1944; $, nth Nov.
1944; §, 2 1 st Jan. 1945. There are also in the same museum from the Itawa
Swamp four specimens of G. media and 1 1 of G. nigripennis, also collected by
myself. Indeed there is a considerable collection of birds in Chicago made
by me in Zambia, as well as in Philadelphia.
In view of this material evidence, the occurrence of G. gallinago in Zambia
is clearly acceptable. Nor does there appear to be reason to question the
records by Tree (1966, Ostrich 37: 186) of birds handled or seen at the Min-
dola Dam, North Kafue Basin, which Benson et al. also, felt to be uncertain.
I did in fact also collect both G. gallinago and nigripennis at Solwezi in January
1 96 1, although I did not retain any specimens.
174
Night Herons in Wallacea
by C. M. N. White
Received 26th September, 197)
Wallacea is used here as a convenient short designation for the Indonesian
islands which lie between the Sunda Shelf (boundary of continental south-
east Asia) and the Sahul Shelf (boundary of continental Australia and New
Guinea). The western boundary is also well known as Wallace's Line.
Darlington (1957: 462-473) has discussed and commended the use of
Wallacea to denote these islands although he did not originate the term
which had been first used nearly thirty years earlier. I use the term only as a
concise geographical designation without implying a zoogeographical region
or subregion.
Nycticorax caledonicus hilli Mathews evidently occurs throughout Wallacea
having been recorded from at least twenty-eight islands. It may be scarce
in the Lesser Sundas west of Timor and Savu for there is only a single
record from Flores, but as it is known to breed in east Java, it seems more
likely to have been overlooked by collectors. Birds from Wallacea agree
with the Australian form hilli, although occasional examples in Celebes show
intergradation towards the darker Philippine manillensis. An abnormally
dark bird from Kema, Minahassa, north Celebes, was named Nycticorax
minahassae Meyer & Wiglesworth but later its authors placed it as a synonym
of manillensis. Amadon (1942: 6) also places it as a synonym of manillensis but
as other examples from north Celebes agree with hilli, it should be more
correctly united with the latter which it would antedate and replace.
Bock (1956) considers that TV. nycticorax and TV. caledonicus form a super-
species. TV. nycticorax has been recorded twice in Wallacea. There is one
record from Flores which is most likely a wanderer from Java or elsewhere in
Malaysia where TV. nycticorax breeds. More remarkable is the occurrence in
the region of Gorontalo in north Celebes. Stresemann (1941: 10) merely
remarks that it very likely breeds at Lake Limbotto but gives no additional
information. Coomans de Ruiter (1948: 80-82) suggests that in fact both
caledonicus and nycticorax are widely sympatric in Celebes and was unable
therefore to identify specifically the birds he saw in south Celebes. Hooger-
werf (1966: 84) also refers to the two species occurring "alongside each
other" in Celebes. The actual situation is however quite different, for TV.
nycticorax is only known from a number of specimens, mostly over a century
old, from the northern peninsula of Celebes. I am very grateful to Dr. G. F.
Mees for kindly giving me details of the material in the Leiden Museum.
There are nine specimens of TV. nycticorax: a male collected by Forsten about
1840: two males, two females and a juvenile female collected in July and
August 1863 by v. Rosenberg at Ayer Pannas, and a juvenile male collected
by the same at Panybie in September: also two unsexed adults, one from
Limbotto in January and another from Menado, undated. The birds obtained
by v. Rosenberg are of most importance, for the same collector also obtained
two juvenile TV. caledonicus in September 1863 at Limbotto and Panybie. Dr.
Mees points out that these juveniles of both species are in spotted plumage
but would have been fully able to fly. There is thus no evidence at all that the
two species of Nycticorax are widely sympatric in Celebes, and only the
specimens collected in 1863 indicate that at that time both species may have
been breeding together in the northern tip of Celebes.
175
Since N. nycticorax and N. caledonicus are both often cited as breeding
in the Philippines, similar sympatry just to the south in north Celebes would
not be surprising. However duPont (1971 : 22) states that iV. nycticorax is only
a migrant from Asia to the Philippines, and Professor D. S. Rabor has kindly
informed me that he, in 39 years of field work in the Philippines, has never
found any evidence that it breeds: he personally believes that it is only a
winter migrant. Thus the suspected sympatric breeding in north Celebes
becomes more remarkable. Confirmation or otherwise of this would be very
desirable, but after a gap of over a century conditions may have changed
and breeding colonies ceased to exist. In the meantime the evidence for this
supposed sympatric breeding is circumstantial only, and relates to 1863.
It is perhaps worth noting that the tropical Asian night heron, Gorsachius
melanolophus •, has not been recorded from Wallacea but that migrants of the
Japanese G. goisagi have been recorded from Celebes and Halmahera.
Hoogerwerf (1966) has described the occurrence of odd examples of N.
caledonicus in a large breeding colony of N. nycticorax in west Java, and
apparent hybridisation with the latter. There was no indication of a mixed
breeding colony. In Borneo N. nycticorax occurs widely and there are a few
records of TV. caledonicus in Sabah in the north but no evidence that they were
breeding. Professor Rabor kindly informs me that he has observed N.
caledonicus throughout the Sulu Archipelago up to the Borneo border so that
Sabah birds might be visitors from this source. Thus all the evidence points
to allopatry between nycticorax and caledonicus being virtually complete. These
extralimital data such as they are only serve to emphasise that if a mixed
breeding colony existed in north Celebes in 1863 it was exceptional.
References :
Amadon, D. 1942. Birds collected during the Whitney South Seas Expedition. 49. Notes on
some non-passerine genera 1. Amer. Mus. Novit. 1175.
Bock, W. J. 1956. A generic review of the family Ardeidae (Aves). Amer. Mus. Novit. 1779-
Coomans de Ruiter, L. 1948. Waarnemingen van Steltloopers (Gressores) in Zuid-Celebes,
in het bijzonder van het blauw-witte reigertje, Notophoyx picata (Gould). Limosa 21:
69-83.
Darlington, P. J. 1957. Zoogeography. New York: Wiley.
duPont, J. 1 97 1. Philippine birds. Greenville, Delaware: Delaware Museum of Natural
History.
Hoogerwerf, A. 1966. The occurrence of Nycticorax caledonicus in Java. Ardea 54: 81-87.
Stresemann, E. 1941. Die vogel von Celebes. 3. Systematik und Biologie. Journ. Ornith. 89:
1-102.
176
CONTRIBUTORS
Contributions are not restricted to members of the Club. They should be
addressed personally to the Editor, C. W. Benson, Department of Zoology,
Downing Street, Cambridge. Contributions are accepted on condition that
sole publication is offered in the first instance to this Bulletin. They should be
type-written, double-spaced, with wide margins, on one side of the paper,
and submitted in duplicate.
References to literature should be in the same format as in the notice to
contributors to the This (see any 1972 number). Considerations similar to
those in the Ibis notice also apply in regard to nomenclature, scientific names
of species and genera, and illustrations including photographs. But illus-
trations including photographs can only normally be considered if the
contributor is willing to pay for the cost of their reproduction.
Contributors introducing a new name or describing a new form should
append nom. nop., sp. nov., subsp. nov. as appropriate. In such a description,
the introduction of the name should be followed by paragraphs for "Des-
cription", "Distribution", "Type", "Measurements of Type", "Material
examined" and further headings as required.
Contributors are entitled to a maximum often free copies of the number of
the Bulletin in which their contribution appears, provided that it exceeds one
page of the Bulletin. Extra copies at cost price can be ordered through the
Editor at the time of submission of the manuscript.
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(vol. 93) and later issues and indices, at 5 op each for vols. 89-92, and at
25P each for pre-1969 issues. Unwanted back numbers returned to the
above address are most gratefully received.
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Other correspondence should be addressed to the Hon. Secretary, R. E. F.
Peal, 24 Creighton Avenue, London N10 1NU.
Published by the BRITISH ORNITHOLOGISTS' CLUB and printed by
The Caxton and Holmesdale Press, 104 London Road, Sevenoaks, Kent.
BULLETIN
of the
BRITISH
ORNITHOLOGISTS' CLUB
EDITED BY
C. W. BENSON
HUGH F. I. ELLIOTT
Volume 94
1974
price: three pounds fifty pence
PREFACE
The quarterly publication and size of the issues which comprise the present volume of the
Bulletin (totalling 176 pp.) have continued on the basis established in 1973. The only minor
change, made for convenience of reference and record, is the resumption of the practice,
last followed in Vol. 89 (1969), of listing the membership of the Club's committee.
The first two issues of 1974 were edited by Mr. C. W. Benson, prior to his retirement
after five years in office, during which he set an example of editorial skill and thoroughness
that would have been quite impossible to follow but for the help and guidance he so freely
gave when handing over. For this, for much subsequent support and, not least, for pre-
paring with the customary and able assistance of Mrs. M. Hawksley, the Scientific Index
for this volume, I am extremely grateful.
As before thanks are due to all who have so kindly advised the editors on papers offered
for publication, including Dr. D. Amadon, Dr. W. R. P. Bourne, Mr. P. R. Colston, Dr.
C. H. Fry, Mr. I. C. J. Galbraith, Mr. D. Goodwin, Mr. M. P. Stuart Irwin, Dr. A. C.
Kemp, Capt. C. R. S. Pitman, Prof. W. H. Thorpe, Mr. R. Wagstaffe and Mr. C. M. N.
White; and likewise to Mr. K. E. Wiltsher, Manager of the Caxton and Holmesdale Press,
and his staff for maintaining standards and punctuality of production of the Bulletin despite
a difficult year for the printing industry.
HUGH F. I. ELLIOTT
COMMITTEE
(1973-74)
Sir Hugh Elliott, Chairman (1971)
Prof. J. H. Elgood, Vice-Chairman (1971)
C. W. Benson, Editor (1969)
R. E. F. Peal, Hon. Secretary (1971)
P. Tate, Hon. Treasurer (1962)
J. H. R. Boswall (1973)
Dr. P. J. K. Burton (1971)
P. Hogg (1972)
C. J. Mead (1971)
(1974-75)
Prof. J. H. Elgood, Chairman (1974)
P. Hogg, Vice-Chairman (1974)
Sir Hugh Elliott, Editor (1974)
R. E. F. Peal, Hon. Secretary (1971)
M. St. J. Sugg, Hon. Treasurer (1974)
J. H. R. Boswall (1973)
Dr. C. J. O. Harrison (1974)
C. J. Mead (1971)
Lt.-Col. J. R. Neighbour (1974)
LIST OF MEMBERS: AMENDMENTS UP TO 31st DECEMBER, 1974
(Compiled by the Hon. Secretary)
New Members
Ashton, P. J., b.sc, Botany Department, Rhodes University, Grahamstown, South Africa.
Dean, W. R. J., Barberspan Nature Reserve, P.O. Barberspan, Transvaal, South Africa.
Dick, Dr. J. A., m.a., m.b., b.chir., 8 Madingley Road, Cambridge CB3 OEE.
Gibbs, A., b.sc, 20 Park Court, Park Road, Hampton Wick, Kingston-upon-Thames,
Surrey KT1 4AV.
Griffin, D., m.a., 51A Palace Road, East Molesey, Surrey.
Hagmann, Dr. J., Kennedy Institute of Rheumatology, Division of Immunology, Bute
Gardens, Hammersmith, London W6 7DW.
Hamel, Rev. P. J., m.a., Chaplaincy at Carleton University, 718 Parkdale Avenue, Ottawa,
Ontario K1Y 1J3, Canada.
Hasegawa, H., M.sc, Department of Zoology-Faculty of Science, Kyoto University,
Sakyo-Ku Kyoto, 606 Japan.
Jensen, J. V. Holtevej 13, DK-8000, Aarhus C, Denmark.
Jones, A. M., 72 Dee Street, Aberdeen AB1 2DS.
Layton, W. A., b.sc, Department of Agriculture, Konedobu, Papua New Guinea.
Limentani, b.sc, b.arch., 10 Kingfisher Green, St. Ives, Huntingdon, Cambridgeshire
PE17 4HS.
McGregor, Dr. I. A., o.b.e., m.r.cp., 210 Hyde End Road, Spencers Wood, near Reading,
Berkshire.
McNeil, Dr. D. A. C, 24 Huntsmoor Road, Ewell, Surrey.
Moller, A. P., Petersborgvej 63, DK-9000, Alborg, Denmark.
Morrison, Dr. R. I. G., Canadian Wildlife Service, 2721 Highway 31, Ottawa, Ontario,
Canada K1A OH3.
Nakata, Y., Kazawu-Cho 5-111, Higasisumiyosi-Ku, Osaka 546, Japan.
Nicholson, M. P., 'Las Rocas', Es Traves, Puerto de Soller, Mallorca, Spain.
Olney, P. J. S., b.sc, Zoological Society of London, Regent's Park, London NW1 4RY.
Oreel, G. J. b.sc, Jozef Israelskade 130-3, Amsterdam-1008, Netherlands.
Parsons, Dr. J., c/o Medical Research Council Project, P.O. Box 1971, Kisumu, Nyanza
Province, Kenya.
Read, W. A., 36 Saltersgate Drive, Birstall, Leicester LE4 3FF.
Sayers, B. C, 164 Chelmer Road, Chelmsford, Essex CM2 6AB.
Tunks, I. D. A., Department of Biological Sciences, Portsmouth Polytechnic, King Henry I
Street, Portsmouth, Hampshire.
Tyler, Dr. S., P.O. Box 10401, Addis Ababa, Ethiopia.
Wolstencroft, J. A., 'Woodside', 10 Clevelands Avenue, Silverdale, Carnforth, Lanca-
shire LA5 0RP.
Resignations
Houston, W. H. Patrick, G. L. A. Sales, V. A. D.
Johnson, F. E. B. Peall, Mrs. O. Searight, R. G.
Lees-Smith, D. T. Russell, J. A. S. Wise, A. J.
Deaths
The Committee much regrets to record the deaths of the following members: —
Colonel the Right Reverend Monsignor F. O. Cave, o.b.e., m.c
Miss J. M. Ferrier
Captain P. R. Foulkes-Roberts, m.c
Miss M. V. Gilbert
Mr. J. M. D. Mackenzie, b.a.
Mr. C. W. Mackworth-Praed, o.b.e., m.a. (Honorary Secretary and Treasurer 1922-
1923 and 1929-1935, Honorary Treasurer 1935-1936, Vice-Chairman 1945-1946,
Chairman 1956-1959).
Professor H. Schouteden
LIST OF AUTHORS AND CONTENTS
Am a don, Dean
Taxonomic notes on the Serpent-eagles of the genus Spilornis
Annual General Meeting (82nd)
Annual General Meeting (82nd — resumed and concluded)
Ash, J. S.
The Boran Cisticola in Ethiopia
Ash, J. S. and Watson, George E.
Locustella naevia in Ethiopia
Auber, L.
The structure of feathers in Chlorophanes purpurascens
Backhurst, G. C.
Buff-breasted sandpiper Tryngites subruficollis in Africa
Benson, C. W.
The type of the Madagascar Cuckoo Cuculus poliocephalus rochii Hartlaub
Another specimen of Neodrepanis hypoxantha
Benson, C. W. and Irwin, M. P. Stuart
The significance of records of the Common Sandpiper breeding in East Africa
Binford, Laurence C. and Zimmerman, Dale A.
Rufous-bellied Heron in Kenya
Blindell, Richard
An address on investigations into waders and wildfowl on the Essex coast and
in the Thames estuary
BOSWALL, JEFFERY
A film on "The private life of the Jackass (Magellanic) Penguin"
Boswall, Jeffery and Freeman, W. P.
A discography of bird sound from the Neotropical Region
Bourne, W. R. P.
The classification of Tristram's Storm-petrel Oceanodroma tristrami Salvin
Britton, Peter L.
Relative Biomass of Ethiopian and Palaearctic Passerines in West Kenya
Habitats
Brooke, R. K.
Buteo tachardus Andrew Smith 1830
The African southern limits of the Steppe Eagle in winter
On the material evidence of Hieraaetus pennatus in Southern Africa
Brown, L. H.
The races of the European Snake Eagle Circaetus gallicus
Bruce, Murray D.
The relationships of Buettikoforella bivittata: some considerations
Bulletin of Zoological Nomenclature
Opinion 999
Cave, F. O.
Notes on birds from the southern Sudan
Clancey, P. A.
On the validity and range of Lamprotornis corruscus mandanus Van Someren
1921
The characters and range of Pogoniulus chrysoconus extoni (Layard), 1871
Colebrook-Robjent, J. F. R. and Osborne, T. O.
High density breeding of the Red-necked falcon Falco chicquera in Zambia
Collins, Charles T. and Tikasingh, Elisha S.
Status of the Great Shearwater in Trinidad, West Indies
Committee, 1973 Report of ..
Conder, P. J.
An address on "Do birdwatchers understand conservation?"
Cooper, J.
The predators of the Jackass Penguin Spheniscus demersus
Davison, G. W. H.
Geographical variation in Lophophorus sclateri
• • 159-
62
41-
-42
.. 129-
131
24-
-26
39-
-40
•• 49
-5 5
176
55
-56
.. 141-
M3
ica 20
-21
. . 101-
102
129
73-76
108-113
59-62
62
152-158
126-128
80-81
128
116-118
113-116
139-141
172-176
96-99
1-2
129
21-24
163-164
Dean, W. R. J.
Bird weights from Angola .. .. .. .. .. .. .. 170-172
Deans, Ian R.
An address on gull ecology in the Thames estuary . . . . . . . . 89
Desfayes, M.
A new subspecies of Accipiter tachiro . . . . . . . . . . . . 69-71
Dillingham, I. H.
Notes on the Lammergeyer Gypaetus barbatus . . . . . . . . . . 72
Donnelly, B. G.
A further record of the White-eyed Gull Larus leucophthalmus from Beira,
Mozambique . . . . . . . . . . . . . . . . . . 62-63
The Lesser Black-backed Gull Larus fuscits in southern and central Africa . . 63-68
Dowsett, R. J.
Geographical variation in iris colour in the bulbul Andropadus milanjensis . . 102-104
Duckworth, Fred
The Whale-headed Stork in Ethiopia . . . . . . . . . . . . 3-4
Elgood, J. H.
Weights and perching habits of birds at Port Moresby, Papua, New Guinea 135-138
Erard, C.
The problem of the Boran Cisticola . . . . . . . . . . . . 26-38
Farkas, T.
On the biology of Monticola imerinus (Hartlaub) . . . . . . . . 165-170
Field, G. D.
Nearctic waders in Sierra Leone — Lesser Golden Plover and Buff-breasted
Sandpiper . . . . . . . . . . . . . . . . . . 76-78
Freeman, W. P.
See Boswall
Frith, C. B.
New observations of migrants and vagrants from Aldabra, Farquhar and
Astove Atolls, Indian Ocean .. .. .. .. .. .. .. 12-19
Fry, C. H.
Vocal mimesis in nestling Greater Honey-guides .. .. .. .. 58-59
Gallagher, M. D.
On the occurrence of the Great White Egret Egretta alba in the Persian Gulf
region .. .. .. .. .. .. .. .. .. .. 122-126
Grimes, L. G.
Duetting in Hyper ger us atrkeps and its taxonomic relationship to Eminia lepida 89-96
Harrison, C. J. O.
A re-examination of material of the extinct marabou stork, Leptoptilos falconeri
with descriptions of some new species
Reduced natal down in some emberizine species
Haverschmidt, F.
Notes on the Gray-breasted Crake Laterallus exilis
The occurrence of the Giant Snipe Gallinago undulata in Surinam . .
Hill, Geoffrey
Observations on a relationship between Crested Guineafowl and Vervet
Monkeys 68-60
Holyoak, D. T.
Cyanoramphus malherbi, is it a colour morph of C. auriceps} . . . . . . 4-9
An address on ornithological exploration and conservation in the Cook
Islands . . . . . . . . . . . . . . . . . . . . 89
Status of the genus Sauropatis (Alcedinidae) .. .. .. .. . .121-122
Undescribed land birds from the Cook Islands, Pacific Ocean .. .. 145-150
Irwin, M. P. Stuart
The Pintail Anas acuta in Rhodesia . . . . . . . . . . . . 56-57
See also Benson
Jefford, T. G.
See Urban
Munteanu, Dan
Further expansion of the Fieldfare in the Rumanian Carpathians . . . . 1 51-15 2
42-49
71-72
2-3
132-134
Oreel, G. J.
Little Gull Larus minutus south of the Sahara . . . . . . . . . . 176
Osborne, T. O.
See COLEBROOK-ROBJENT
Parkes, Kenneth C.
Geographical variation in the Rufous-tailed Foliage gleaner Philydor rufi-
caudatus, with notes on plumages .. .. .. .. .. .. 11 8-1 21
Payne, Robert B.
Species limits and variation of the New World Green Herons Butorides
virescens and Striated Herons B. striatus . . . . . . . . . . 81-88
Sassoon, S.
Finfoot Podica senegalensis in Ethiopia . . . . . . .... . . 176
Scott, Sir Peter
An illustrated talk on individuality in birds . . . . . . . . . . 1
Sinclair, J. C.
Larus minutus in Angola . . . . . . . . . . . . . . . . 57
Fish offal scavengers off Luanda . . . . . . . . . . . . 58
Special General Meeting .. .. .. .. .. .. .. 132
Thorpe, W. H.
An illustrated talk on individuality in bird voices . . . . . . . . 41
TlKASINGH, ELISHA S.
See Collins
Turner, D. A.
Cape Grass Owl in Ethiopia .. .. .. .. .. .. .. 38-39
Urban, Emil K. and Jefford, T. G.
The status of the cormorants Phalacrocorax carbo lucidus and Phalacrocorax
carbo patricki . . .. .. .. .. .. .. .. .. 104-107
Walters, Michael P.
The eggs of the Golden Conure Aratinga guarouba . . . . . . . . 71
Watson, George E.
See Ash
Wattel, J.
An illustrated talk on moult in relation to life cycle in some non-passerines . . 1
White, C. M. N.
Three water birds of Wallacea . . . . . . . . . . . . . . 9-1 1
The Little Egret Egret ta garget ta in Wallacea . . . . . . . . . . 78-79
Butastur and Buteo east of Wallace's Line . . . . . . . . . . 99-100
Some questionable records of Celebes birds . . . . . . . . . . 144-145
WlNTERBOTTOM, J. M.
Turtle Dove Streptopelia turtur in South West Africa . . . . . . . . 19
Zimmerman, Dale A.
See Binford
INDEX TO SCIENTIFIC NAMES
(Compiled by C. W. Benson with the assistance of Mrs. M. Hawksley)
All generic and specific names (of birds only) are indexed. Subspecific names are included
only if new and are also indexed in bold print under the generic and the specific names.
abbotti, Spilornis 162
Accipiter fasciatus 137-138
— tachiro 69-70
Accipiter tachiro croizati, subsp. nov. 70
Acrocephalus palustris 112
— schoenobaenus 112
— scirpaceus 40, 111
— vaughani 149-150
Acrocephalus vaughani kaoko, subsp.
nov. 150
Acrocephalus vaughani kerearako,
subsp. nov. 149
acuta, Anas 56
adelaidae, Platycercus 6
aethiopicus, Threskiornis 22
alba, Calidris 77
— , Egretta 79, 122-125
— , Motacilla 16
— , Tyto 38
albescens, Morphninus 155
albicollis, Ficedula 1 1 1
albifrons, Platysteira 171
alboscapularis, Malurus 137-138
albus, Corvus 13, 173-174
Alca alle 128
— impennis 128
aldabranus, Dicrurus 13, 15
alle, Alca 128
amethystina, Chalcomitra 117
Anabazaenops immaculatus 118
analoga, Meliphaga 136
Anas acuta 56
— platyrhynchos 18
— querquedula 56
andrewsi, Fregata 145
Andropadus milanjensis 102-104
— tephrolaemus 102
— virens 171
angolensis, Gypohierax 58
anguitimitens, Eurocephalus 157
Anhinga rufa 12
ansorgei, Cossypha 171
Anthus trivialis 16
Apalis flavida 171
— pulchra 95
apivorus, Pernis 60
Aplonis cantoroides 136-138
Apus apus 15
apus, Apus 1 5
Aquila nipalensis 62, 152
— pomarina 152
— wahlbergi 155-156
aquila, Fregata 145
Aratinga guarouba 71
Ardea cinerea 58, 123
— novaehollandiae 9
Ardeola ralloides 101
ardeola, Dromas 14, 19
Arenaria interpres 18
Argala arvenensis 42
— falconeri 42
Artamus leucorhynchus 136-13!
arvenensis, Argala 42
asturinus, Spilornis 162
atiu, Halcyon 148
atricapilla, Sylvia 40, 111-112
atriceps, Hypergerus 89-96
— , Moho 89
auratus, Oriolus 171
auriceps, Cyanoramphus 4-8
ayresi, Sarothrura 39
badiceps, Eremomela 171
bairdii, Prinia 171
Balaeniceps rex 3
Barbatula extoni 139
barbatus, Gypaetus 72
Bathmocercus cerviniventris 95
Batis minulla 171
beaudouini, Circaetus 126-128
bensoni, Monticola 167
biarmicus, Falco 172
Bias musicus 171
bicolor, Nigrita 172
bivittata, Buettikoforella 80
bocagei, Telophorus 171
bodessa, Cisticola 32-38
borin, Sylvia 109, 111-113
Bowdleria punctata 80
brachyptera, Cisticola 171
bradfieldi, Tockus 171
brevipes, Monticola 165
brevirostris, Phalacrocorax 107
Bucorvus leadbeateri 171
Buettikoforella bivittata 80
bulliens, Cisticola 171
bulocki, Merops 58-59
Butaetes lessonii 1 5 5
Butastur indicus 100
— liventer 99-100
Buteo buteo 59-61, 100
desertorum 60, 100
— lagopus 60
— oreophilus 59-60
Buteo rufofuscus 59
— tachardus 59-61
buteo, Buteo 59—61, 100
Butorides rufiventris 101
— striatus 81-88, 101
— sundevalli 81, 87
— virescens 81-88
Bycanistes sharpii 171
Cacomantis variolosus 137-138
Cairina moschata 18
caledonicus, Nycticorax 136
Calidris alba 77
— canutus 77
— melanotos 76
— minuta 14
Camaroptera fasciolata 95
— superciliaris 171
canicapilla, Nigrita 172
caniceps, Lonchura 136-138
canorus, Cuculus 15, 59
cantoroides, Aplonis 136-138
canutus, Calidris 77
capensis, Smithornis 94
— , Tyto 38
capistrata, Nesocharis 117
capistratum, Pellorneum 80
caprata, Saxicola 137
carbo, Phalacrocorax 104-107
carteri, Eremiornis 80
carunculatus, Phalacrocorax 106-107
castanea, Dyaphorophyia 171
castaneothorax, Lonchura 137-138
castanotus, Colius 171
Catharacta maccormicki 21
cayana, Dacnis 51—55
Centropus phasianinus 136-138
Cercomela familiaris 117
cerviniventris, Bathmocercus 95
— , Chlamydera 137-138
Ceryle maxima 171
— rudis 171
Chalcomitra amethystina 117
chalconotus, Phalacrocorax 107
Charadrius forbesi 116
— hiaticula 77
cheela, Spilornis 159-163
chelicuti, Halcyon 171
chicquera, Falco 172-175
chiniana, Cisticola 25-38
Chlamydera cerviniventris 1 3 7-1 3 8
Chlidonias nigra 57-58
Chlorocichla falkensteini 171
Chlorophanes purpurascens 49-54
— spiza 51-55
chloropygia, Nectarinia 171
chrysaetos, Falco 153
chrysoconus, Pogoniulus 139-14 1
Cichlornis grosvenori 80
Ciconia 43, 48
Ciconia, ciconia 21
— episcopus io-ii
— stormi 10- 11
ciconia, Ciconia 21
cinerea, Ardea 58, 123
Circaetus beaudouini 126-128
— gallicus 126-128
— pectoralis 126-128
cirrocephalus, Larus 63, 66
Cisticola bodessa 32-38
Cisticola bodessa kaffensis, subsp. nov.
32
Cisticola brachyptera 171
— ■ bulliens 171
— chiniana 25-38
— exilis 136-138
— natalensis 31
— subruficapilla 32
Cly tospiza monteiri 117
coelebs, Fringilla 25, 27
Colius castanotus 171
Collocalia leucophaea 146-147
Collocalia sawtelli, sp. nov. 146-147
Collocalia vanikoriensis 136
Colluricincla harmonica 136-138
collurio, Lanius 40
collybita, Phylloscopus 40
Columba livia 17, 157
columbarius, Falco 19, 174
Columbigallina passerina 133
comitatus, Pedilorhynchus 171
communis, Sylvia 27, 40
concolor, Falco 13
Coracina novaehollandiae 136-138
— papuensis 136-138
— pectoralis 171
coronatus, Spizaetus 155
corruscus, Lamprotornis 11 3-1 15
coruscans, Neodrepanis 141-143
Corvus albus 13, 173-174
— orru 136-138
Cosmetornis vexillarius 170
Cossypha ansorgei 171
Cracticus mentalis 135-138
crassirostris, Hemiparra 101
crestatus, Eudyptes 23
Crinifer leucogaster 157
— personata 157
cristatus, Lanius 1 1 1-1 1 2
croizati, Accipiter 70
cruentatum, Dicaeum 144
crumeniferus, Leptoptilos 43
Cryptociconia indica, gen. et. sp. nov. 44,
48-49
Cuculus canorus 15,59
— poliocephalus 15, 55-56
— rochii 56
curruca, Sylvia 40
cuvieri, Falco 175
Cyanerpes cyaneus 51-55
cyaneus, Cyanerpes 5 1-5 5
Cyanoramphus auriceps 4-8
— intermedius 6
— malherbi 4-8
— novaezelandiae 7-8
Cyanosylvia svecica 168
Dacelo gaudichaud 135-136
— leachii 135, 137-138
Dacnis cayana 51-55
Delichon urbica 21
demersus, Spheniscus 21-22
desertorum, Buteo 60, 100
diadematum, Tricholaema 157
Dicaeum cruentatum 144
— hosei 144-145
— nehrkorni 144
Dicrurus aldabranus 13, 15
Dissoura neglecta 10
dominica, Pluvialis 77-78
dominicanus, Larus 22, 58, 63-64, 66
Dromas ardeola 14, 19
Drymocataphus 80
dubius, Hieraaetus 155
— , Leptoptilos 43-44, 47, 49
Dumetia hyperythra 80
dumonti, Mina 137-138
Dyaphorophyia castanea 171
edouardi, Guttera 68
Egretta alba 79, 122-125
— garzetta 58, 78-79, 122
— gularis 122
— intermedia 79, 124-125
elegans, Platycercus 6
eleonorae, Falco 13
elgini, Spilornis 159-163
Elminia longicauda 171
Eminia lepida 89-96
episcopus, Ciconia 10-11
epops, Upupa 40
Eremiornis carted 80
Eremomela badiceps 171
erythrocephalus, Trachyphonus 157
erythrocercus, Philydor 120
Erythrocnus 101
erythrops, Neocrex 3
Estrilda melpoda 172
Eudynamys scolopacea 144
Eudyptes crestatus 23
euophrys, Philydor 120
Eurocephalus anguitimitens 157
Eurystomus glaucurus 16
— orientalis 137
exilis, Cisticola 136-138
— , Ixobrychus 73
— , Laterallus 2-3
explorator, Monticola 167, 169
extoni, Barbatula 139
falcinellus, Plegadis 11
Falco biarmicus 172
— chicquera 172-175
— chrysaetos 153
— columbarius 19, 174
— concolor 13
— cuvieri 175
— eleonorae 13
— longipennis 175
— newtoni 13, 16
— pennatus 152
— peregrinus 12-13
— subbuteo 175
— tachardus 60
falconeri, Argala 42
— , Leptoptilos 42-45, 49
falkensteini, Chlorocichla 171
familiaris, Cercomela 117
fanovanae, Newtonia 143
fasciatus, Accipiter 137-138
— Tockus 171
fasciolata, Camaroptera 95
Ficedula albicollis m
flava, Motacilla 101, 111-113
flaveola, Sicalis 71
flaveolus, Platycercus 6
flavescens, Meliphaga 135-138
flavida, Apalis 171
flavigaster, Hyliota 171
— •, Microeca 136-138
flavipes, Tringa 76
flaviventer, Porzana 2
forbesi, Charadrius 116
Foudia madagascariensis 18
fraseri, Stizorhina 171
Fregata andrewsi 145
— ■ aquila 145
— ■ minor 145
Fringilla coelebs 25, 27
fuliginosa, Nectarinia 171
fusconota, Nigrita 172
fuscus, Larus 18, 58, 63-68
galatea, Tanysiptera 136
Galerida modesta 95
gallicus, Circaetus 126-128
Gallinago gallinago 20, 133
— hardwicki 134
— macrodactyla 20
— nigripennis 20
— stenura 20
— undulata 132-134
gallinago, Gallinago 20, 133
garzetta, Egretta 58, 78-79, 122
gaudichaud, Dacelo 135-136
Geopelia humeralis 137-138
— striata 18, 136-138
giganteus, Macronectes 27
gilvus, Mimus 133
glareola, Tringa 78
glaucurus, Eurystomus 16
goodwini, Ptilinopus 145
Graminicola 80
gravis, Puffinus 96
grosvenori, Cichlornis 80
guarouba, Aratinga 71
gularis, Egretta 122
Guttera edouardi 68
gutturalis, Irania 40
— , Neocichla 171
— , Phalacrocorax 106
Gypaetus barbatus 72
Gypohierax angolensis 58
haematodus, Trichoglossus 136-138
Haematopus ostralegus 77
Halcyon chelicuti 171
— leucocephala 171
— macleayii 136-137
— miyakoensis 122
Halcyon ruficollaris, sp. nov. 147
Halcyon sancta 122, 136-138
— senegalensis 171
Halcyon tuta atiu, subsp. nov. 148
Halcyon tuta mauke, subsp. nov. 148-149
Haliaeetus vocifer 173
hardwicki, Gallinago 134
harmonica, Colluricincla 136-138
Hemiparra crassirostris 101
heuglini, Ploceus 117
hiaticula, Charadrius 77
Hieraaetus dubius 155
— pennatus 60, 152-158
— spilogaster 155
Himantopus himantopus 101
himantopus, Himantopus 101
Hippolais pallida 40
Hirundo lagopoda 60
— rustica 16, 19, 111-113
— tahitica 136-138
hosei, Dicaeum 144-145
humeralis, Geopelia 137-138
— Myospiza 133
huttoni, Phalacrocorax 107
Hyliota flavigaster 171
Hypergerus atriceps 89-96
hyperythra, Dumetia 80
Hypochera nigeriae 1 1 7
hypoleucos, Tringa 14, 20, 77
hypoxantha, Neodrepanis 1 41-143
imerinus, Monticola 165-170
immaculatus, Anabazaenops 118
impejanus, Lophophorus 163-164
impennis, Alca 128
indica, Cryptociconia 44, 48-49
Indicator indicator 58-59
— minor 58
— variegatus 58
indicator, Indicator 58-59
indicus, Butastur 100
insignis, Prodotiscus 171
intermedia, Egretta 79, 124-125
intermedius, Cyanoramphus 6
interpres, Arenaria 18
Irania gutturalis 40
Ixobrychus exilis 73
jacarina, Volatinia 2
jardineii, Turdoides 171
javanica, Rhipidura 144
javanicus, Leptoptilos 44-45
jugularis, Nectarinia 137-138
Jynx torquilla 40
kaffensis, Cisticola 32
kaoko, Acrocephalus 150
kerearako, Acrocephalus 149
kinabaluensis, Spilornis 160-163
klossi, Spilornis 162
lacrymosum, Tricholaema 157
Lagonosticta nigricollis 117
— rubricata 117
— rufopicta 117
— senegala 117
lagopoda, Hirundo 60
lagopus, Buteo 60
Lalage sueurii 136-138
Lamprotornis corruscus 11 3-1 15
Lanioturdus torquatus 61
Lanius collurio 40
— cristatus 111-112
— nubicus 40
— souzae 171
Larus cirrocephalus 63, 66
— dominicanus 22, 58, 63-64, 66
— fuscus 18, 58, 63-68
— leucophthalmus 62, 66
— minutus 57, 176
Laterallus exilis 2-3
— melanophaius 3
— viridis 3
leachii, Dacelo 135, 137-138
leadbeateri, Bucorvus 171
lepida, Eminia 89-96
Leptoptilos crumeniferus 43
— dubius 43-44, 47, 49
— falconeri 42-45, 49
— javanicus 44-45
Leptoptilos richae, sp. nov. 43-44, 47, 49
Leptoptilos siwalicensis, sp. nov. 44, 46,
49
lessonii, Butaetes 155
leucocephala, Halcyon 171
leucogaster, Crinifer 157
leucolaima, Pogoniulus 171
leucophaea, Collocalia 146-147
— , Microeca 136-138
leucophrys, Rhipidura 136-138
leucophthalmus, Larus 62, 66
leucopygius, Turdoides 61, 171
leucorhynchus, Artamus 136-138
lhuysii, Lophophorus 163-164
liventer, Butastur 99-100
livia, Columba 17, 157
Locustella naevia 39-40
Lonchura caniceps 136-138
— castaneothorax 137-138
longicauda, Elminia 171
longipennis, Falco 175
Lophophorus impejanus 163-164
— lhuysii 163-164
— sclateri 163-164
Lophophorus sclateri orientalis, subsp.
nov. 164
Luscinia megarhynchos 40
— svecica 40, 165, 168, 170
maccormicki, Catharacta 21
macleayii, Halcyon 136-137
Macrocephalon maleo 144
macrodactyla, Gallinago 20
— , Oceanodroma 100
Macronectes giganteus 21
madagascariensis, Foudia 18
— Phyllastrephus 143
magellanicus, Spheniscus 23, 129
magna, Sturnella 133
Malaconotus 6-7
maleo, Macrocephalon 144
malherbi, Cyanoramphus 4-8
Malurus 80
— alboscapularis 137-138
Mandingoa nitidula 117
mariae, Megalurulus 80
markhami, Oceanodroma 100
matsudairae, Oceanodroma 100
mauke, Halcyon 148
maxima, Ceryle 171
maximus, Thalasseus 58
Megalurulus mariae 80
megarhynchos, Luscinia 40
melania, Oceanodroma 100
melanoleucos, Phalacrocorax 106-107
melanophaius, Laterallus 3
melanopis, Schistochlamys 133
melanopsis, Monarcha 136
melanotos, Calidris 76
Meliphaga analoga 136
— flavescens 135-138
melpoda, Estrilda 172
mentalis, Cracticus 135-138
— , Sphenoeacus 95
Merops bulocki 58-59
— ornatus 136-137
— superciliosus 15
meyeri, Poicephalus 170
Microeca flavigaster 136-138
— leucophaea 136-138
migrans, Milvus 138
milanjensis, Andropadus 102-104
milvoides, Spizaetus 154
Milvus 158
— migrans 138
Mimus gilvus 133
Mina dumonti 137-138
minimus, Spilornis 162
minor, Fregata 145
— , Indicator 58
minulla, Batis 171
minuta, Calidris 14
minutus, Larus 57, 176
miyakoensis, Halcyon 122
modesta, Galerida 95
Moho atriceps 95
Monarcha melanopsis 136
monteiri, Clytospiza 117
Monticola bensoni 167
— brevipes 165
— explorator 167, 169
— imerinus 165-170
— saxatilis 167
— sharpei 166-167, I ^9
— solitarius 167
monticola, Oenanthe 169
Morphninus albescens 1 5 5
moschata, Cairina 18
Motacilla alba 16
— flava 101, 111-113
Muscicapa striata 17, 111-112
musicus, Bias 171
Myospiza humeralis 133
nabouroup, Onychognathus 61
naevia, Locustella 39-40
Napothera 80
natalensis, Cisticola 31
nebularia, Tringa 21, 77
Nectarinia chloropygia 171
— fuliginosa 171
— jugularis 137-138
— olivacea 95, 171
— seimundi 171
— superba 171
— verticalis 171
neglecta, Dissoura 10
nehrkorni, Dicaeum 144
Neocichla gutturalis 171
Neocrex erythrops 3
Neodrepanis coruscans 1 41-143
— hypoxantha 1 41-143
Nesocharis capistrata 117
newtoni, Falco 13, 16
Newtonia fanovanae 143
nigeriae, Hypochera 117
nigerrimus, Ploceus 172
nigra, Chlidonias 57-58
nigricollis, Lagonosticta 117
nigripennis, Gallinago 20
Nigrita bicolot 172
— canicapilla 172
— fusconota 172
nipalensis, Aquila 62, 152
nisoria, Sylvia 40
nitidula, Mandingoa 117
novaeguineae, Philemon 136-138
novaehollandiae, Ardea 9
— , Coracina 136-138
novaezelandiae, Cyanoramphus 7-
nubicus, Lanius 40
Numenius phaeopus 76
Nycticorax caledonjcus 136
Oceanodroma macrodactyla 100
— markhami 100
— matsudairae 100
— melania 100
— tristrami 100
Oenanthe 168
— monticola 169
— oenanthe 16
— pleschanka 40
oenanthe, Oenanthe 16
olivacea, Nectarinia 95, 171
— ■, Tiaris 72
Onychognathus nabouroup 61
oreophilus, Buteo 59-60
orientalis, Eurystomus 137
— , Lophophorus 164
Oriolus auratus 171
— oriolus 17
— szalayi 136-138
oriolus, Oriolus 17
ornatus, Merops 136-137
orru, Corvus 136-138
Ortygocichla rubiginosa 80
ostralegus, Haematopus 77
Otus scops 40
pallida, Hippolais 40
pallidirostris, Tockus 171
paludicola, Riparia 112
palustris, Acrocephalus 112
papuensis, Coracina 136-138
parasiticus, Stercorarius 58
passerina, Columbigallina 133
pectoralis, Circaetus 126-128
Pedilorhynchus comitatus 171
Pellorneum capistratum 80
peltata, Platysteira 171
pennatus, Falco 152
— Hieraaetus 60, 152-158
peregrinus, Falco 12-13
Pernis apivorus 60
personata, Crinifer 157
Petrochelidon rufigula 171
phaeopus, Numenius 76
Phalacrocorax carbo 104-107
— carunculatus 106-107
Phalacrocorax chalconotus 107
— gutturalis 106
— huttoni 107
— melanoleucos 106-107
phasianinus, Centropus 136-138
Philemon novaeguineae 136-138
philippensis, Rallus 137-138
Philydor erythrocercus 120
— euophrys 120
— ruficaudatus 11 8-1 21
— subflavescens 119
Phoeniculus purpureus 171
Phoenicurus phoenicurus 40, 112
phoenicurus, Phoenicurus 40, 112
Pholidornis rushiae 172
Phyllastrephus madagascariensis 143
— tenebrosus 143
— zosterops 143
Phylloscopus collybita 40
— sibilatrix 17
— trochilus 16-17, 4°> m-112
picturata, Streptopelia 14
pilaris, Turdus 151
Pinguinus 128
Platycercus 7
— adelaidae 6
— elegans 6
— flaveolus 6
platyrhynchos, Anas 18
Platysteira albifrons 171
— peltata 171
Plautus 128
Plegadis falcinellus 1 1
pleschanka, Oenanthe 40
pleurostictus, Thryothorus 73
Plocepasser rufoscapulatus 171
Ploceus heuglini 117
— nigerrimus 172
— vitellinus 117
Plotus 128
plumata, Prionops 171
Pluvialis dominica 77-78
— squatarola 13, 77
Podica senegalensis 170, 176
Poicephalus meyeri 170
— rueppellii 170
Pogoniulus chrysoconus 1 39-141
— leucolaima 171
— pusillus 139
poliocephalus, Cuculus 15, 55-56
pomarina, Aquila 152
Porzana flavi venter 2
Prinia bairdii 171
Prionops plumata 171
— retzii 171
Prodotiscus insignis 171
Ptilinopus rarotongensis goodwini,
subsp. nov. 145-146
Puffinus gravis 96
— puffinus 98
puffinus, Puffinus 98
pulchra, Apalis 95
punctata, Bowdleria 80
purpurascens, Chlorophanes 49-54
purpureus, Phoeniculus 171
pusillus, Pogoniulus 139
querquedula, Anas 56
ralloides, Ardeola 101
Rallus philippensis 137-138
rarotongensis, Ptilinopus 145-146
retzii, Prionops 171
rex, Balaeniceps 3
Rhipidura javanica 144
— leucophrys 136-138
richae, Leptoptilos 43-44, 47, 49
Riparia paludicola 112
— riparia 111-113
riparia, Riparia 111-113
rochii, Cuculus 56
rubetra, Saxicola 111-112
rubiginosa, Ortygocichla 80
rubricata, Lagonosticta 117
rudis, Ceryle 171
rueppellii, Poicephalus 170
rufa, Anhinga 12
rufescens, Sylvietta 171
ruficapilla, Spermophaga 172
ruficaudatus, Philydor 11 8-1 21
ruficollaris, Halcyon 147
rufigula, Petrochelidon 171
rufipectus, Spilornis 162
rufiventris, Butorides 101
rufofuscus, Buteo 59
rufopicta, Lagonosticta 1 1 7
rufoscapulatus, Plocepasser 171
rushiae, Pholidornis 172
rustica, Hirundo 16, 19, 111-113
Salpornis spilonota 171
sancta, Halcyon 122, 136-138
Sarothrura ayresi 39
saturata, Scolopax 20
Sauropatis 121
sawtelli, Collocalia 146-147
saxatilis, Monticola 167
Saxicola 168
— caprata 137
— rubetra 111-112
— torquata 40
Schistochlamys melanopis 133
schoenobaenus, Acrocephalus 1 1 2
scirpaceus, Acrocephalus 40, 111
sclateri, Lophophorus 163-164
scolopacea, Eudynamys 144
Scolopax saturata 20
scops, Otus 40
seimundi, Nectarinia 171
senegala, Lagonosticta 1 1 7
senegalensis, Halcyon 171
— , Podica 170, 176
— , Streptopelia 157
sharpei, Monticola 166-167, l< ^9
sharpii, Bycanistes 171
sibilatrix, Phylloscopus 17
Sicalis flaveola 71
sipora, Spilornis 162
siwalicensis, Leptoptilos 44, 46, 49
Smithornis capensis 94
solitaria, Tringa 76
solitarius, Monticola 167
souzae, Lanius 171
Spermophaga ruficapilla 172
Sphecotheres vieilloti 137-138
Spheniscus demersus 21-22
— magellanicus 23, 129
Sphenoeacus 80
— mentalis 95
spilogaster, Hieraaetus 155
spilonota, Salpornis 171
spilonotus, Spilornis 160, 162
Spilornis abbotti 162
— asturinus 162
— cheela 159-163
— elgini 159-163
— kinabaluensis 160-163
— klossi 162
— minimus 162
— rufipectus 162
— sipora 162
— spilonotus 160, 162
spiza, Chlorophanes 51-55
Spizaetus coronatus 155
— milvoides 154
squatarola, Pluvialis 13, 77
Stachyris 80
stenura, Gallinago 20
Stercorarius parasiticus 58
stormi, Ciconia 10-11
Stipiturus 80
Stizorhina fraseri 171
Streptopelia picturata 14
— senegalensis 157
— turtur 14-15, 19
striata, Geopelia 18, 136-138
— , Muscicapa 17, 111-112
striatus, Butorides 81-88, 101
Sturnella magna 133
subbuteo, Falco 175
subflavescens, Philydor 119
subruficapilla, Cisticola 32
subruficollis, Tryngites 77, 176
sueurii, Lalage 136-138
sundevalli, Butorides 81, 87
superba, Nectarinia 171
superciliaris, Camaroptera 171
superciliosus, Merops 15
svecica, Cyanosylvia 168
— , Luscinia 40, 165, 168, 170
Sylvia atricapilla 40, 111-112
— borin 109, 111-113
— communis 27, 40
— curruca 40
— nisoria 40
Sylvietta rufescens 171
— virens 95, 171
szalayi, Oriolus 136-138
tachardus, Buteo 59-61
— , Falco 60
tachiro, Accipiter 69-70
tahitica, Hirundo 136-138
Tanagra violacea 71
Tanysiptera galatea 136
Telacanthura ussheri 1 5
Telmatodytes 80
Telophorus bocagei 171
tenebrosus, Phyllastrephus 143
tephrolaemus, Andropadus 102
Tersina viridis 71
Thalasseus maximus 58
Threskiornis aethiopicus 22
Thripias xantholophus 171
Thryothorus pleurostictus 73
Tiaris olivacea 72
Tockus bradfieldi 171
— fasciatus 171
— pallidirostris 171
torquata, Saxicola 40
torquatus, Lanioturdus 61
torquilla, Jynx 40
totanus, Tringa 77
Trachyphonus erythrocephalus 157
Trichoglossus haematodus 136-138
Tricholaema diadematum 157
— lacrymosum 157
Tringa flavipes 76
— glareola 78
— hypoleucos 14, 20, 77
— nebularia 21, 77
Tringa solitaria 76
— totanus 77
tristrami, Oceanodroma 100
trivialis, Anthus 16
trochilus, Phylloscopus 16-17, 4°> III_
112
Tryngites subruficollis 77, 176
Turdoides jardineii 171
— leucopygius 61, 171
Turdus pilaris 151
turtur, Streptopelia 14-15, 19
tuta, Halcyon 148-149
Tyto alba 38
— capensis 38
undulata, Gallinago 132-134
Upupa epops 40
urbica, Delichon 21
vanikoriensis, Collocalia 136
variegatus, Indicator 58
variolosus, Cacomantis 137-138
vaughani, Acrocephalus 149-150
verticalis, Nectarinia 171
vexillarius, Cosmetornis 170
vieilloti, Sphecotheres 137-138
violacea, Tanagra 71
virens, Andropadus 171
— Sylvietta 95, 171
virescens, Butorides 81-88
viridis, Laterallus 3
— , Tersina 71
vitellinus, Ploceus 1 1 7
vocifer, Haliaeetus 173
Volatinia jacarina 2
wahlbergi, Aquila 155-156
xantholophus, Thripias 171
zosterops, Phyllastrephus 143
Corrigenda
15, line 33: Insert inverted commas after 'specimen'
p. 49, line 5 : i dubius\ not 'dubia
p. 80, line 20: i mariae\ not i mariei''
p. 95, line 23: i Sphenoeacus\ not 'Sphenoaecus'
p. 95, line 25 : 'Camaroptera, not l Cameropterd
p. 136, line 9: < phasianinus\ not ' 'phasianus''
p. 170, penultimate line: l rueppellii\ not 'ruppe//i'
p. 171, line 1: l castanotus\ not ' ' castonotus'
p. 171, line 43: i ret^ii\ not 'w/^/V
p. 173, table: ' Haliaeetus\ not 'Haliaaetus^
The Caxton & Holmesdak Press, Sevenoaks
Bulletin of the
British Ornithologists' Club
Edited by
C. W. BENSON
Volume 94 No. 1
March 1 974
Committee
Sir Hugh Elliott, Bt., o.b.e. {Chairman)
J. H. Elgood (Vice-Chairman)
C. W. Benson, o.b.e. (Editor)
R. E. F. Peal (Hon. Secretary)
P. Tate (Hon. Treasurer)
J. H. R. Boswall
Dr. P. J. K. Burton
P. Hogg
C. J. Mead
Bulletin of the
BRITISH ORNITHOLOGISTS' CLUB
1
Vol. 94 No. 1 Published: 20 March, 1974
L " * Ap %l97d\
The six hundred and eighty-fifth meeting of the Club was held at the Cafe
Royal, 68 Regent Street, London, W.i, on Tuesday 27th November 1973
at 7 p.m.
Chairman: Sir Hugh Elliott, BlTo^Te., present 16 members and 6 guests.
The speaker was Dr. Jan Wattel who addressed the Club on Moult in
relation to life cycle in some non-passerines and illustrated his talk with
slides.
The six hundred and eighty-sixth meeting of the Club was held at the Cafe
Royal, 68 Regent Street, London, W.i, on Tuesday 15th January 1974 at
7 p.m.
Chairman: Sir Hugh Elliott, Bt., o.b.e., present 26 members and 18 guests.
The speaker was Sir Peter Scott, c.b.e., d.s.c, who addressed the Club on
Individuality in birds and illustrated his talk with slides of Bewick's Swans
Cygnus bewickii and other wildfowl.
Report of the Committee for 1973
Six meetings of the Club were held in 1973, all at the Cafe Royal. The number
of members and guests attending was 176, compared with 191 in 1972.
Despite a prospective increase of charges in 1974, the advantages of the Cafe
Royal as a venue were considered, after due investigation, to justify a con-
tinuation of present arrangements, except possibly for holding one Buffet
Supper meeting (probably in July).
Leaflets, which described the advantages of membership of the Club and,
for those ineligible to join as members, of subscription to the Bulletin^ were
enclosed in the July 1973 number of Ibis and have contributed to the satis-
factory numbers of 30 new members and five new non-member subscribers
to the Bulletin in 1973. The Committee regrets to report the death of two
members, Mr. R. P. Borrett and Mr. K. D. Smith, and there were three
resignations, whilst the membership of seven members was terminated under
Rule 4. There was thus an increase in membership of 18. An increased num-
ber of members and of non-member subscribers is important for the financial
stability of the Club. The leaflet is continuing to bring in some new members
and subscribers but it is hoped that members will recommend membership
of the Club or subscription to the Bulletin (£3 • 50 p. a. for non-members) to
friends or institutions who may possibly be interested.
Four numbers of the Bulletin were published in 1973, with a total of 176
pages. Arrangements have now been made for authors of papers exceeding
one page in the Bulletin to obtain separates of them and 16 separates will be
supplied free in respect of each such paper, commencing with the March
1974 number of the Bulletin. Mr. C. W. Benson ends his term of office as
Editor at the Annual General Meeting. He is unable to stand for re-election
owing to the pressure of other work over the next year. Sir Hugh Elliott has
agreed to be nominated as Editor but hopes to be able to resign in favour of
Mr. Benson if the latter feels able to accept nomination in 1975.
Copies of the Accounts will be available zt the Annual General Meeting
and will be published subsequently in the Bulletin. It is not possible to comment
here in detail on the financial position, as this Report must go to press before
the Accounts are available. It is expected, however, that there will be a deficit on
the year, as sales of back-numbers of the Bulletin have not reached the
abnormally high figure of 1972. Difficulty has been caused in recent years by
members who have not paid subscriptions due or who have paid only at a
lower rate than has become current (£1 • 50 p. a.). Steps are being taken to re-
duce the loss from this cause and it is satisfactory to report that in several cases
arrears have already been paid. In view of the Club's need for support, to meet
costs which still tend to increase despite the saving made by quarterly publica-
tion of the Bulletin, it is much to be hoped that the process will continue.
Notes on the Grey-breasted Crake Laterallus exilis
by F. Haver schmidt
Received jth November, 1973
In Surinam Laterallus exilis is not uncommon in densely overgrown marshes
where it is usually only seen when flushed and flying off with dangling legs low
over the vegetation to alight within a short distance (Haverschmidt 1968).
Hence it is difficult to collect. In these rather wet places it is often in company
with the still smaller Yellow-breasted Crake Por^ana flaviventer.
In December 1972 I established that it also inhabits a totally different dry
habitat. On the outskirts of rapidly expanding Paramaribo there is a large
house building project on the former plantations Tourtonne and Ma Retraite.
Streets had been built, and between them were open plots, well drained and dry,
of area about 200 by 100 metres and thickly overgrown with tall grass to a
height of about one metre. On 23 December a mowing machine was cutting
grass, starting at the outside and steadily moving inward. When only a narrow
strip of tall grass was left, a crowd of small boys followed, running on each side
of the machine. Repeatedly a small crake rose from the remaining grass,
fluttered a short distance to plane down a few metres farther on the open ground
when it was immediately covered by a bunch of the boys and easily captured.
Very few birds escaped, and within five minutes I had four birds. On 27
December two machines were at work in the same area, five more birds being
taken, and on 30 December I obtained another one. The number of crakes in
these extensive grass fields must indeed have been very large. No other species
of crake was observed or captured. The only other bird inhabiting these fields
in large numbers was the Blue-black Grassquit Volatinia jacarina.
Of this series of 10 specimens, five were males and five females, and nine
were in adult plumage. Four males and four females were in breeding condition,
the males with greatly enlarged testes and females with enlarged ovaries. One (
female (27 December) held a fully developed, although unshelled, egg in the
oviduct. So it was obvious that these specimens had been captured in the breed-
ing season. Previously, on 4 March 1961, I had obtained two nestlings still
completely in black down. In plumage, the immature specimen (male) lacks
the chestnut on the nape, hindneck and mantle. It is uniform plain brown on the
upperparts, except for the grey crown. Two other immature specimens which
I collected on 22 and 29 January 1961 are similar. There is great variation in
the pale barring on the wing coverts. In only one male is it prominent, and on
four females it is very faint. In the others — one male and four females, including
the immature — the wing coverts are plain brown with no barring at all.
Although generally similar in colour, the males are easily distinguishable by
the somewhat darker grey on the breast, contrasting more strongly with the
white throat than in the females.
The iris was red with an orange eyelid, the legs ochre yellow. In the adults
the upper mandible was horn brown except for the lower part of the base,
which was greenish yellow (as in illuminating paint), extending to the nostrils.
The lower mandible was also the same greenish yellow. In immatures the upper
mandible was wholly horn brown, and the greenish yellow was only faintly
apparent on the lower mandible.
As eight out of the nine birds in adult plumage were in breeding condition,
it is somewhat strange that one of these nine (adult female, 27 December) was
moulting all its primaries, all still in sheath, so that at this time it would have
been unable to fly. It is clear that the wing moult of Laterallus exilis is syn-
chronous as in L. melanophaius but different from L. viridis (Stresemann &
Stresemann 1966). This moulting female was the only adult in the series which
was not in breeding condition.
Some of the small South American crakes may be wanderers, as was estab-
lished by Beebe (1947) for Neocrex erjthrops in Venezuela. Two records from
Surinam suggest that this may also apply to L. exilis, since I received two
females (2 July i960 and 13 February 1962) that had been captured at night at
lighted houses in Paramaribo.
My 10 specimens weighed: 5^ 32-4 - 37-7 (34*6) g, 5$$ 28-9 - 39-0
(34-0) g. The heaviest female was the one which had an unshelled egg in the
oviduct.
References :
Beebe, W. 1947. Avian migration at Rancho Grande in north central Venezuela. Zoologica
32: 166.
Haverschmidt, F. 1968. Birds of Surinam. Edinburgh & London: Oliver & Boyd.
Stresemann, E. & Stresemann, V. 1966. Die mauser der vogel. Journ. Ornith. 107, sonderheft:
141-155.
The Whale-headed Stork in Ethiopia
by Fred Duckworth
Received 3rd July, 1973
Mr. T. Mattanovich reported seeing the Whale-headed Stork Balaeniceps rex
Gould along the Baro River in western Ethiopia, west of Gambela, both in the
rains (May-October) and in the dry season (November-April). This was duly
recorded by Urban (1967) and again by Urban & Brown (1971).
While on safari in the Gambela District in the first fortnight of April 1973
I saw Whale-headed Storks as follows : — firstly, at the Ubela Waterhole
freshwater reedbeds, 7°3o'N, 34°o3'E, one bird; secondly, a few days later, in
similar habitat just north of Tedo, 7°5 5'N, 34°03'E, two pairs together. On
both occasions the birds were very shy, allowing an approach no closer than
75 metres. But both times I was able to obtain a photograph and so positively
confirm the occurrence of this species in Ethiopia. Local information is that the
Whale-headed Stork is resident in Gambela, although there is not yet a
breeding record.
References :
Urban, Emil K. 1967. Possible occurrence of the Whale-headed Stork in Ethiopia. Journ,
E. Afr. Nat. Hist. Soc. 26(2): 87-88.
Urban, Emil K. & Brown, Leslie H. 1971. A checklist of the birds of Ethiopia. Addis Ababa:
Haile Sellassie I Univ. Press.
[The author of the above note is a Regional Wild Life Warden in Ethiopia Copies of the
two photographs which he obtained, unquestionably of Balaeniceps rex, have been deposited
in the University Museum of Zoology, Cambridge. — Ed.].
Cyanoramphus malherbi, is it a colour morph
of C. auriceps ?
by D. T. Holy oak
Received 22nd October, iyj)
The Orange-fronted Parakeet Cyanoramphus malherbi of New Zealand has been
treated as a valid species by almost every authority since it was first described
by De Souance in 1857. Examination of museum specimens and a search of the
literature have convinced me that the taxonomic status of this bird needs to be
reassessed. This paper discusses the available evidence bearing on the relation-
ship of C. malherbi and the Yellow-fronted Parakeet C. auriceps auriceps,
leading to the conclusion that C. malherbi is probably a colour morph of C. a.
auriceps, although direct proof is lacking.
Si%e and Structure: Oliver (1955), Falla et al. (1970) and Harrison (1970)
quote early accounts which describe C. malherbi as slightly smaller than C. a.
auriceps, with a smaller, more slender, bill. The ranges of measurements quoted
by Oliver seem to confirm this size difference, but they do not agree with my
own measurements (cf. Table I). Table I gives data suggesting that bill-length
measurements of both forms are bimodally distributed. The two peaks pre-
sumably correspond to a sexual dimorphism in size, but few accurately sexed
specimens are available to test this hypothesis. Wing-length measurements
may also be bimodally distributed, but with much more overlap than those for
bill-length. Statistical testing shows a high probabliity of both sets of measure-
ments being representative of the same population (Table I).
TABLE 1
Measurements of Cyanoramphus malherbi and C. auriceps auriceps. Because few accurately
sexed specimens are available all measurements for each form are listed together. X = mean,
n = sample size, s == standard deviation.
Bill length (mm)
9 10 ii 12 13 14 15 16 X n s X±2s
C. malherbi
01543510 12-4 19 1-39 9-6-15-2
C. a. auriceps
1 1 10 1 3 11 3 2 12-7 32 1-82 9-1-16-3
Wing length (mm)
100- 102- 104- 106- 108- no- 112- 114- 116-
101 103 105 107 109 in 113 115 117 X n s X ± 2s
C. malherbi
3 2 5 1 5 1 o 1 o 107 18 3-67 99-114
C. a. auriceps
3 4 5 7 4 4 2 2 4 108 35 4-82 98-117
4
Fewer comparative measurements of the lengths of tail and tarsus, and of
the depth and width of the bill, likewise show no significant differences between
C. malherbi and C. a. auriceps. Nor are there consistent differences in the shape
and structure of the bill, legs or feet.
There are neither consistent differences nor large average differences between
the two forms in the shape and relative lengths of the individual remiges and
rectrices, nor in the rest of the plumage.
Colouration: The usual colouration of the plumage and bare parts of each
form may be compared as follows (checked with ten specimens of each form) :
C. a. auriceps
C. malherbi
Forehead
scarlet
orange
Crown
golden-yellow
lemon-yellow
Lores
scarlet above, yellow-
orange above, light green
green below
below
Chin
light yellow-green
light green
Ear-coverts
yellow-green
green
Sides of neck, nape,
yellow-green
green
mantle, back, scapulars,
upper tail-coverts
Rump
yellow-green with
green with small patch
small patch of
of orange-tipped
scarlet-tipped
feathers at side
feathers at side
Throat, breast, belly,
light yellow-green
light green
thighs, under tail-
coverts
Exposed parts of
yellow-green
green
greater, median and
lesser wing-coverts
Exposed parts of primary-
deep blue
deep blue
coverts and
feathers of alula
Primaries
blackish-brown with
blackish-brown with blue
Secondaries
Lesser underwing-
coverts
Greater underwing-
coverts
Rectrices
Bill
Cere
Iris
Legs and feet
blue on outer webs of
outer primaries, tur-
quoise on outer webs
of inner primaries
green outer webs
light green
blackish-brown
middle pair mainly
green above, others
turquoise and blue
above
Blackish-grey with
large blue-grey area
in middle of upper
mandible
blackish-grey
orange-yellow
dark grey
on outer webs of most
primaries, a little tur-
quoise on outer webs of
innermost
blue-green outer webs on
outermost secondaries, green
outer webs on others
light blue-green
blackish-brown
middle pair mainly green
above, others blue with
little turquoise
blackish-grey with
large blue-grey area
in middle of upper
mandible
blackish-grey
unknown
dark grey
Yellow plumage colouration in parrots is caused by a little-known "floures-
cent" pigment, by carotenoids, or by both in some species (Volker 1937,
Auber 1957). Blue colouration is caused by the Tyndall effect of scattering of
short wave-lengths of light by gas filled vacuoles in the keratin of the feather
(Auber 1957, Dyck 1971). Green colouration is caused by the combination of
the Tyndall scattering effect with one or both of the yellow pigments. Red
colouration is apparently due to carotenoid pigments (Auber 1957, Fox &
Vevers i960, Dyck 1971).
I have examined museum skins of C. a. auriceps and C. malherbi in ultra-violet
light and could detect flourescence from neither the yellow nor green feathers
(the flourescent yellow pigment retains its ability to flouresce in old museum
skins: Holyoak unpublished). Hence, from Volker's studies it is apparent that
the yellow plumage pigment in these two forms is a carotenoid.
When the descriptions of the colouration of C. a. auriceps and C. malherbi
given above are analysed in terms of this account of pigmentation it is clear
that differences in carotenoid pigmentation are sufficient to account for all of
the colour differences. To summarise: —
Forehead, Rump
patches
Crown
Green areas of
plumage
Remiges and Rectrices
Heavier carotenoid
pigmentation
(C. a. auriceps)
scarlet
golden-yellow
have strong yellow
tint
more extensive green
areas (structural
blue -j- yellow
carotenoid = green)
Lighter carotenoid
pigmentation
(C malherbi)
orange
lemon- yellow
lack strong yellow
tint
some green areas of
C. a. auriceps
replaced by blue
(green — yellow
carotenoid = blue)
There is widespread evidence that birds judged to be closely related from
other characters have shown rapid evolutionary change in carotenoid pig-
mentation. For example, Cain (1955) described the apparent hybridisation of
Platycercus elegans (predominantly red plumage) and P. flaveolus (predominantly
yellow plumage), with P. "adelaidae" (predominantly orange plumage) as a
stable intermediate form. Holyoak (1973) gives data suggesting that the genera
Platycercus and Cyanoramphus are closely related.
Additional evidence that the extent and colour of avian carotenoid pigmen-
tation can be under simple genetic control is provided by the striking poly-
morphisms of some shrikes of the genus Malaconotus (Moreau & Southern 1958,
Hall, Moreau & Galbraith 1966). The apparently simple genetic control of
carotenoid pigmentation may in part be due to the inability of birds, like other
animals, to synthesise carotenoids (cf. Vevers 1964).
Harrison (1970) states that there are 59 specimens of C. malherbi in museums
of the world, whereas there must be about 300 of C. a. auriceps. When the small
total number of specimens of each form is considered, it appears remarkable
that so many with unusual colouration have been described, even allowing for
the special value that the Victorian collectors attached to "varieties". C. W.
Benson {in litt.) has described a specimen which has the dull green body colour
of C. malherbi with the red forehead of C. a. auriceps (i8/Psi/i9/a/6 in the
Cambridge Museum Collection). Another intermediate specimen was described
as Cyanoramphus intermedius Reichenow; from the description of this bird in
Salvadori (1891) it appears to have been a typical specimen of C. a. auriceps \
except that it had the lemon-yellow crown colour of C. malherbi; I have been
unable to locate the specimen. Harrison (1970) mentions a specimen of C.
malherbi that Souance described as having a barely distinguishable orange fore-
head band. Three specimens of C. a. auriceps in the Collection of the British
6
Museum (Natural History) at Tring show plumage differences from the 17
normal specimens of this form in the Collection: — 1847. 1.8. 40 has scarlet tips
to many feathers in middle of rump; 1897. 11. 10. 644 has scarlet marks on the
golden-yellow crown feathers; 195 3.62.2 has a few orange marks on the breast
feathers.
Thus, although the colour differences between C. a. auriceps and C. malherbi
seem large at first sight, comparatively small differences in pigmentation are
sufficient to account for all of them. These differences are of a kind that seem
to be under simple genetic control in the closely related genus, Platycercus and
are of a similar kind to the phenotypic differences in a classic instance of avian
polymorphism in the genus Malaconotus. In addition there are two specimens
showing some characters of one form with some of the other, and several
others showing individual variation in the extent or kind of carotenoid pig-
mentation.
Range and Status: C. a. auriceps is known from the North and South Islands
of New Zealand, Three Kings, Hen Island, Barrier Islands, Kapiti, Solander
Islands, Stewart Island and outlying islets, and the Auckland Islands (Fleming
1953, Kinsky et al. 1970). C. a. forbesi, a much larger representative form of C.
auriceps, occurs on the Chatham Islands. C. malherbi is definitely known only
from South Island and Stewart Island; old records from Hen Island and Little
Barrier Island lack sufficient supporting detail to be considered fully acceptable
and old records from North Island are poorly substantiated (Fleming 1953,
Kinsky et al. 1970, Harrison 1970). C. malherbi has not been reported from any
locality on South Island at which C. a. auriceps is unknown, and the older
unconfirmed reports of this form are all from islands where C. a. auriceps is
common.
Oliver (1955) and others described a decrease in the status of C. a. auriceps
and C. novae^elandiae on North and South Islands that occurred through the
first part of this century. Both species remained common on the smaller
outlying islands. Turbott (1967), and the annual lists of New Zealand bird
observations published in Notomis, suggest that C. a. auriceps has regained
much lost ground in the last few decades, so that it is now quite common in the
larger forest tracts; C. novae^elandiae has remained rare.
Harrison (1970) summarises all of the known records of C. malherbi, of which
there were numerous reports in the nineteenth century, but only seven this
century, in 1904, 1913, 1928, then a gap until 1949, 1955, 1963 and 1965.
i Generally, it appears that the decrease of C. malherbi approximately coincided
with those of C. a. auriceps and C. novae^elandiae, although it appears to have
always been rarer than those forms. The recent records of C. malherbi could
reflect a recent increase in its status, as well as increased observation.
Habitat, Composition of Flocks: Early accounts suggested that C. malherbi was
restricted to montane scrub and forest at high elevations, but Harrison (1970)
summarises many later records which show that it occurs mainly at lower
elevations than this, in similar forests to those occupied by C. a. auriceps.
Most C. malherbi seen in the field have been associated with parties or flocks
of C. a. auriceps. Haast probably saw more C. malherbi in the field than any other
observer; he stated {fide Oliver 1955), that "this species and the Yellow-fronted
Parakeet occur always together, but in some localities the first and in others the
second is predominant". Some of the recent reports of C. malherbi were at sites
where C. a. auriceps was seen at about the same time (Anon. 1949, Breen 1956,
I 959)-
Food: Reischek (1885) said that the food of C. malherbi consists of berries
and seeds, and Harrison (1970) quotes a label on a specimen in the British
Museum (Natural History) stating "stomach small grubs". The food of C. a.
auriceps consists of a wide variety of vegetable matter, ranging from fruit and
seeds, to buds, leaves and the nectar and pollen of flowers. Insects have not
been recorded in the diet of this form, but they are probably taken in small
quantities as they are by many other platycercine parrots.
Behaviour and Nesting: Hardly anything has been recorded of the behaviour of
C. malherbi, and very little of that of C. a. auriceps. Falla et al. (1966) state that
the voice of C. malherbi apparently bears a close resemblance to those of C.
auriceps and C. novae^elandiae, but details are not available in the literature.
Apart from a clutch of three eggs said to be of C. malherbi from Mt. Peel
which are now in the Canterbury Museum, New Zealand, no nest of this form
has been recorded in the wild. The literature appears to lack any reference to
either pure pairs of C. malherbi or mixed pairs with C. a. auriceps.
C. malherbi has only once been bred in captivity. From the account of Prest-
wich quoted by Harrison (1970), it would appear that a pair kept by De
Laurier produced a brood of four young in France in 1883. De Laurier also kept
C. auriceps at that time, but unfortunately there is no indication of whether or
not they were in separate cages from the breeding pair of C. malherbi ', nor are
there descriptions of the nestlings, all of which might have died before they
were fully feathered from the meagre information given.
Discussion of the Status ofC. malherbi: The information given above makes it
reasonable to hypothesise that C. malherbi is a colour morph of C. a. auriceps
rather than the distinct species it has been widely assumed to be.
The differences between these forms in size and bill structure that are often
quoted appear to be based on the study of too few specimens, so that what
appears to be a sexual size difference may have been mistaken for a species
difference. Other structural differences appear to be lacking, and the colour
differences can be explained in terms of small changes in carotenoid pigmen-
tation that are probably under simple genetic control. Evidence that C. malherbi
is only known from areas in which C. auriceps occurred, and that C. malherbi
"always" occurred in flocks of C. auriceps when it was more common, support
the colour-morph hypothesis. Insufficient is known of the behaviour, vocali-
sations and nesting of both forms for these kinds of information to be used in
assessing their status.
Evidence that bill size and structure are similar in the two forms suggests
they have similar feeding ecology. As most C. malherbi have apparently been
seen in mixed flocks with C. a. auriceps in typical habitats for the latter, it is
difficult to see how the ecological segregation required by the Volterra-Gause
Principle could be maintained, although such segregation could occur only
at times of food shortage. Congeneric species of island birds often show larger
differences in bill size than continental species (Lack 1971), and with parrots
generally, closely related sympatric species occurring together in the same
habitats usually differ markedly in size, particularly bill size. These ecological
reasons tend to suggest that the two forms are conspecific.
Observations on the occurrence of mixed pairs or of offspring differing from
the parents are necessary to test the colour morph hypothesis, and these are
not available. As C. malherbi is now rare it seems unlikely that field-workers in
New Zealand will be able to fill these gaps in the knowledge available, lthough
this is possible if a sighting of this form is followed up by intensive observation
and nest-hunting. In the meantime, it is probably best to treat C. malherbi as a
synonym of C. a. auriceps on the basis of the inferential evidence available.
Acknowledgements : I am grateful to the staff of the Sub-department of Orni-
thology, British Museum (Natural History) for allowing access to specimens in j
their care, and particularly to Dr. D. W. Snow for help in consulting the
literature. Mr. P. J. Morgan made a number of useful suggestions and helped
me to examine specimens in his care at the Liverpool Museum. Mr. C. W.
Benson was kind enough to examine and measure specimens in the Cambridge
Museum for me and made useful suggestions. Mrs. M. LeCroy kindly measured
specimens in the American Museum of Natural History for me, and Dr. C. J. O.
Harrison made helpful comments on my manuscript.
References :
Anon. 1949. Bird life in Collin's Valley, Wakatipu. A^. Z. Bird Notes 3: 100.
Auber, L. 1957. The distribution of structural colours and unusual pigments in the class
Aves. Ibis 99: 463-476.
Breen, M. J. 1956. In "Classified summarised notes". Notornis 6: 207.
— 1959- In "Classified summarised notes". Notornis 8: 74.
Cain, A. J. 1955. A revision of Trichoglossus haematodus and of the Australian platycercine
parrots. Ibis 97: 432-479.
Dyck, J. 1 97 1. Structural and spectral reflectance of green and blue feathers of the Rose-
faced Lovebird Agapornis roseicollis. K. danske Vidensk. Selsk. Biol. Skr. 18: 1-67.
Falla, R. A., Sibson, R. B. & Turbott, E. G. 1970. A field guide to the birds of New Zealand.
2nd ed. London: Collins.
Fleming, C A.etal.195}. Checklist of New Zealand birds. Wellington, N. Z.: A. H. & A. W.
Reed for Orn. Soc. of New Zealand.
Fox, H. M. & Vevers, G. i960. The nature of animal colours. London: Sidgwick & Jackson.
Hall, B. P., Moreau, R. E. & Galbraith, I. C. J. 1966. Polymorphism and parallelism in the
African bush shrikes of the genus Malaconotus (includes Chlorophoneus). Ibis 108:
161-182.
Harrison, M. 1970. The Orange-fronted Parakeet Cyanoramphus malherbi. Notornis 17: 1 1 5—
I25 '
Holyoak, D. T. 1973. Comments on taxonomy and relationships in the parrot subfamilies
Nestorinae, Loriinae and Platycercinae. Emu 73: 157-176.
Kinsky, F. C. et al. 1970. Annotated checklist of the birds of New Zealand. Wellington, N. Z.:
A. H. & A. W. Reed for Orn. Soc. N. Z.
I Lack, D. 1 971. Ecological isolation in birds. Oxford & Edinburgh: Blackwell.
Moreau, R. E. & Southern, H. N. 1958. Geographical variation and polymorphism in
Chlorophoneus shrikes. Proc. %ool. Soc. London 130: 301-328.
Oliver, W. R. B. New Zealand birds. 2nd. ed. Wellington, N. Z.: A. H. & A. W. Reed.
Reischek, A. 1885. On the habits of New Zealand birds. Trans. N. Z. Inst. 18: 98.
Salvadori, T. 1891. Catalogue of the Psittaci, or Parrots, in the Collection of the British
Museum. Cat. Bds. Brit. Mus. 20. London.
Turbott, E. G. (ed.) 1967. Butter's birds of New Zealand (reprint of Buller 1888, 2nd. ed.).
London: MacDonald.
Vevers, G. 1964. Article "Colour" in Thompson, A. L. (ed.), New Diet. Birds. London:
Nelson.
Volker, O. 1937. Uber fluoreszierende, gelbe federpigmente bei papageien, eine neue
klasse von federfarbstoffen. fourn. Ornith. 85: 136-146.
Three water birds of Wallacea
by C. M. N. White
Received 14th November, 197)
This note provides new information on three species of heron, stork and ibis
which occur in Wallacea.
Ardea novaehollandiae
This common Australian heron has been recorded from a number of islands
in southern Wallacea: Lombok, Sumbawa, Flores, Sumba, Savu, Timor,
Roma, Babar, Tenimber Islands, Kei Islands, Manggoer, Manawoka, Goram
and Madu. There is also a record from Kema, north Celebes. Amadon (1942:
2-3) reviewed geographical variation in this species, finding birds from Walla-
cea identical with those of Australia. He did not mention that birds in Wallacea
are probably non-breeding migrants from Australia.
Rensch (1931: 502) found the species common in Lombok, Sumbawa and
Flores from March to July, in swamps and rice paddies near the coast and
flighting to roosts in the evening. Specimens collected were not in breeding
condition, and from March to early May were in wing moult. Other species of
heron breed in these islands from April to July.
I have assembled the dates of 23 specimens collected in Wallacea. All fall
between March and October apart from one from Timor in February. In north
Australia breeding takes place from December to February. All the data from
Wallacea seems to indicate that there is marked movement into a well defined
part of Wallacea of migrants after breeding, and that this species must be added
to the number of Australian birds which regularly winter in Wallacea.
Ciconia episcopus
This stork occurs in Wallacea in Celebes, Muna and Buton: and in Lombok,
Sumbawa and Flores. They have sometimes been considered a subspecies
distinct from the nominate Asiatic C. episcopus, under the name C. e. neglecta,
but there has been uncertainty about this. When Finsch described Dissoura
neglecta in 1904 he did not indicate the material upon which the name was based
but merely gave the range as "Java, Sumbawa, Lombok, Celebes, Philippines".
However in Notes from the Leyden Museum, 16 July, 1905 : 152 he states that
he had five specimens from Java, a similar number from Celebes and one from
Sumbawa, and that the Philippine birds were "probably" the same as these.
Riley (1924: 28) recorded four specimens from Celebes which differed from
birds from Burma, Thailand and the Philippines in having the distal two-thirds
of the bill red with basal third black instead of having the bill dusky with only
the tip and a line along the culmen red. He adopted the name neglecta for them,
but pointed out that the exact type locality of that form was not known, and
that in any case Philippine birds were like nominate episcopus.
Chasen (193 5 : 53) places neglecta as a synonym of episcopus, giving its type
locality as Java, and stating in a footnote "We cannot separate D. e. neglecta".
Stresemann (1941 : 2) confirms the bill colour described by Riley in respect of
two more birds from Celebes and adds that one from Lombok agrees with them.
I am very grateful to Dr. G. F. Mees for information on questions discussed
here. All the birds examined by Finsch are still in the Leiden Museum except
for one from Java. None of the specimens was designated as the type, so that
all are syntypes. All show the extensively red bills described by Riley and
Stresemann. Dr. Mees himself observed birds in west Java in January 1949, and
noted at the time that their bills were dark red brown, black at base and along
ridges. D. e. neglecta it appears can be separated from nominate episcopus by the
colour of the bill, and birds from Java agree with those of Wallacea. The dark
bill of the nominate form, apart from descriptions in the literature, is shown
well in a coloured photograph in Kahl (1971 : plate V). Chasen erred in giving
the type locality of neglecta as Java, for he gives no indication that he was in-
tending to designate a restricted type locality.
It seems possible that nominate episcopus is separated from neglecta by a wide
geographical gap. For the Malay peninsula Chasen gives episcopus as only
occurring in the north: further south he gives only Perak where C. stormi is
recorded, considered by Chasen as a separate species. C. stormi occurs in
Borneo and is represented from east Sumatra (U.S. National Museum) and from
North Pagi, West Sumatran Islands (Mus. Comp. Zool., Harvard). Chasen
quotes Sumatra as part of the range of C. episcopus but this needs verification
from any specimens that may exist. Mr. I. C. J. Galbraith kindly informs me
that there are none in the British Museum (Nat. Hist.), Tring. Unless it can be
shown that nominate episcopus does occur in Sumatra, stormi is interposed
between it and neglecta.
Plegadis falcinellus
The status of the glossy ibis in Wallacea is obscure. It is evidently well known
in Celebes where it was first collected at Makassar in^i828. There are series
of adult and immature birds in the Leiden Museum, but Dr. G. F. Mees kindly
informs me that there are no juveniles or any trace of evidence that it breeds.
Raven collected eight adults and immatures in March 191 7, at Rano Lindoe in
north-central Celebes. Coomans de Ruiter (195 1 : 307) recorded large flights in
V-formation along the coast in south-west Celebes in February with a flock of
over 100 on 5 April.
If these many glossy ibises do not breed in Celebes it is difficult to imagine
their origin. The species is unrecorded in Thailand or southern Vietnam: it
is also almost unknown in Malaysia, with a single record from south Borneo
in 185 1, and a breeding population in west Java. In the Philippines it is re-
corded from Luzon and Mindanao and Delacour & Mayr (1946: 32) state
"resident and breeds locally in Mindanao". It is difficult to believe that large
numbers of birds move from either west Java or Mindanao to Celebes.
There are no records from southern Wallacea, and van Bemmel (1948) does
not list it from the Moluccas. Actually Meyer & Wiglesworth (1898, ii: 803)
do mention a specimen from Ternate in north Moluccas, but perhaps van
Bemmel discarded this as one of the many plumage trade records bearing this
as a fictitious locality. However Mrs. M. LeCroy has kindly informed me that
in the American Museum of Natural History there is a female, AMNH 531287,
Halmahera, October, 1884, Bruijn collectors. It seems therefore that these
Moluccan records justify adding the species to the list of birds known from
these islands. They are likely to be vagrants from Australia and there are few
records from New Guinea, which are also suspected of being birds moving
from Australia. These records from both Moluccas serve to emphasise the
curious position in Celebes, even though they provide additional localities for
Wallacea.
References :
Amadon, D. 1942. Birds collected during the Whitney South Sea Expedition. XLIX. Notes
on some non-passerine genera. Amer. Mus. Novit. 1175.
Chasen, F. N. 1935. A hand list of Malaysian birds. Bull. Raffles Mus. 11.
Coomans de Ruiter, L. 195 1. Vogels van het dal van de Bodjo-rivier (Zuid-Celebes).
Ardea 39: 261-318.
Delacour, J. & Mayr, E. 1946. Birds of the Philippines. New York: Macmillan.
Kahl, M. P. 1 971. Social behaviour and taxonominc relationships of the storks. The Living
Bird 10: 1 51-170.
Meyer, A. B. & Wiglesworth, L. W. 1898. The birds of Celebes and the neighbouring islands.
Berlin: Friedlander.
Rensch, B. 1931. Die vogel von Lombok, Sumbawa und Flores. Mitt. Zool. Mus. Berlin 17:
451-637.
Riley, J. H. 1924. A collection of birds from north and north-central Celebes. Proc. U.S.
Nat. Mus. 64(16): 1-118.
Stresemann, E. 1941. Die vogel von Celebes. Journ. Ornith. 89: 1-102.
Van Bemmel, A. C. V. 1948. A faunal list of the birds of the Moluccan Islands. Treubia 19:
323-402.
New observations of migrants and vagrants
for Aldabra, Farquhar and Astove Atolls, Indian Ocean
by C. B. Frith
Received 2nd November, 1973
Introduction
Since 1966 Aldabra Atoll (see maps in Stoddart 1971 for location) has been
the subject of considerable scientific attention, brought about partly by a
threat to use it as a military airfield and as a site for a major broadcasting
station (Stoddart 1968). In 1971 The Royal Society of London opened a
Research Station south of the Old Settlement on West Island on Aldabra,
after plans for the above projects were halted as a result of economic and
political pressures. I was engaged at this station as a member of the scientific
staff from 9 April 1972 to 2 April 1973, and it was from there that I carried
out field work for the present paper. My main concern was with the land
birds and it must be stressed that notes given here on shore birds are the
result of purely incidental observations.
The majority of records are from direct observation. Whenever possible,
any doubtful birds were photographed for confirmation of identification or
specimens were collected. Benson & Penny (1971) dealt with all the species
of land birds recorded for Aldabra, and where applicable I have made com-
parative comments. A few records are added for Astove and Farquhar Atolls
made during a day stop en route for the Seychelles from Aldabra.
The scientific nomenclature followed is that of Forbes-Watson (1972)
being the most recent for the Malagasy Region. The following symbols are
used after the species name : —
* new record for atoll.
j- specimen obtained.
% photograph obtained.
Specimens and photographs are in the British Museum (Natural History),
Tring.
Systematic List
Anhinga rufa, African Darter *l
On 16 July 1972 a Seychelles labourer reported a "new bird with a very
long dagger-like beak". A bird fitting this description was observed by a
number of the Station staff on 29 July and identified as an African Darter.
The bird was subsequently seen at least five times, the last date being 16
September 1972. All sightings took place around the Research Station and
West Channels. The bird was in fine condition and at low water was seen
perched on small rocks projecting a foot or two above the water surface on
the open reef flat. It was only observed swimming once, in the deep water at
Passe Femme where the tidal flow was considerable. Attempts to find the bird
after 16 September failed. It could have been either A. r. rufa of Africa or
vulsini of Madagascar.
Falco peregrinus, Peregrine Falcon *{
Two individuals were seen, both in the vicinity of the Research Station.
The first observed on 12 December 1972 was in immature plumage and the
photographs obtained leave no doubt as to its identification. On 23 March
1973 a second was observed, perched in sparse Casuarina woodland, and a
good view was had for about two minutes. This individual was in fine adult
plumage. Probably these were both the palaearctic F. p. calidus, recorded
from Africa (White 1965 : 61) and from Mauritius (Benson & Penny 1971 :
515).
Falco eleonorae , Eleonora's Falcon *f
Two specimens were collected near the Old Settlement, on 1 and 17
November respectively. Both were light phase female birds, with un-
developed oocytes and fully ossified skulls. The first was in very worn
plumage with no sign of moult; it had a fresh dead weight of 273 g and a
wing length of 309 mm. The second was less worn with no sign of active
moult but with new primaries and secondaries. The fresh dead weight was
270 g and wing length 323 mm. The gizzards of the specimens were empty
save for the remains of one, unidentifiable, ant.
A live specimen was found on 1 December 1972 in very worn plumage, it
was extremely thin and one eye had recently been damaged badly. This was
possibly due to Pied Crows Corvus albus which were seen to attack other
large falcons (see below). This bird was kept in captivity for eight weeks and
then set free. Benson & Penny (1971 : 515) give an almost certain sight record
for 4 March (1968), but the above two are the first entirely certain records.
Others are included below.
Falco spp.
A considerable number of falcons were seen between June 1972 and March
1973, most of them appearing in November and December. All were larger
than the local kestrel, F. neirtoni, and mostly of the appearance of F. eleonorae.
Those of which I had poor views could possibly have been F. eleonorae or F.
concolor. All observations are given below and were made near or within the
Old Settlement save the first: —
16 June 1972: Cinq Cases Camp, South Island. Brief view of large, dark
falcon.
31 October 1972: With little doubt a single F. eleonorae seen in flight.
16 November 1972 : Poor views of single bird but appeared smaller than
F. eleonorae and not as rufous on ventral side. Snatching insects in flight,
once seen to hold insect in feet and feed in flight. Hovered for short
periods once or twice.
21 November 1972 : Three falcons soaring and gliding. Two identical to
specimens of F. eleonorae collected and one much darker. One was being
mobbed by a Pied Crow.
25 November 1972 : A dark falcon flying very high and being chased by
four Pied Crows.
27 November 1972: A lone falcon seen at very close quarters, perched
six m high in Casuarina tree. I have no doubt this bird was a dark phase
F. eleonorae.
2 December 1972: A dark falcon was seen twice during the day, prob-
ably the same individual. On one occasion it was being severely mobbed
by a kestrel, F. newtoni, and on the other by a Pied Crow and an Aldabra
Drongo Dicrurus aldabranus.
Clearly the above falcons, including F. peregrinus and F. eleonorae, were part
of a migration movement passing over Aldabra mostly during November
and December. These birds appeared to stay a very short time on the atoll,
if at all, possibly as a result of mobbing by resident birds or lack of suitable
food.
Pluvialis squatarola, Grey Plover
Single or small groups of up to four birds were seen near the Research
13
Station during every month except July, August, October and January. In
addition, approximately a dozen were observed feeding on sand flats between
He Michel and South Island on 17 June 1972, and four at Dune Jean-Louis on
5 March 1973. A single bird in both December and March was in breeding
plumage, otherwise all individuals were in non-breeding dress. Probably this
species can be found on Aldabra in every month, as can be inferred from the
records quoted by Penny (1971: 552).
Tringa hypokucos, Common Sandpiper
Only one individual seen, feeding on the sand bar off-shore from
Research Station, 9 May 1972. This does not appear to be at all a common
species on Aldabra.
Calidris minuta, Little Stint
Noted on two occasions: — 3 October 1972: Twelve in a mixed flock
(including Turnstone, Greater Sand Plover, Whimbrel and Crab Plover) off-
shore from Research Station; 5 March 1973: Approximately 200, forming
major part of a mixed flock (including Grey Plover, Greater Sand Plover and
Crab Plover) on beach and reef flat at Dune Jean-Louis, South Island.
Curlew Sandpipers were also present both times in small numbers for
comparison. The records in Penny (1 971 : 5 5 3) do not indicate that the Little
Stint can be normally numerous on Aldabra.
Dromas ardeola, Crab Plover
Seen throughout the year, in flocks usually consisting of 20 to 50 indi-
viduals. Conspicuously larger flocks, approximate numbers per flock, were
as follows : —
17 May 1972: 1,000 birds on lies Moustique.
25 October 1972 : 450 at Dune Jean-Louis.
27 October 1972: 250 flew past Research Station from south to north.
5 February 1973 : 450 flew past Research Station from south to north.
7 February 1973 : 400 flew past Research Station from south to north.
The larger flocks observed flying northward are worthy of note. It is true
some flocks were seen flying south past the Station but these were far smaller
in number and loosely organised compared with the larger ones, and were
probably feeding birds. The northward bound flocks, however, were in
tight formation, flying low and fast in a direct line. They were not seen to
alight or break up whilst within sight of the Station, after which point they
would shortly have reached the north-west tip of the atoll and the open sea.
A projected line of their flight path would lead directly to the coast of
Somalia, where the species is known to breed from March to October (Archer
6 Godman 1937 : 447). The October flock seen flying north would, however,
arrive at the end of the breeding season. In any case there seems to be little
doubt that Aldabra plays an important role in the movements of this species.
Streptopelia turtur^ Turtle" Dove *f
A single bird appeared on 4 December 1972, and seen feeding on rice with a
number of Madagascar Turtle Doves S. picturatahy a chicken pen. D. Good-
win of the British Museum (Nat. Hist.) examined the specimen and provided
the following notes: — "I think this bird is of the nominate race, S. t. turtur.
It agrees well in size with samples of this form and the differences in coloura-
tion are, I think, due to wear and weathering. Although it is darker than
specimens (in the British Museum collections) of S. t. turtur (beacuse of wear,
I think), one would have expected it to be less dark were it referable to the
paler (and smaller) S. t. arenicola, the larger, reddish population probably
14
breeding in the Yarkland regions (Goodwin 1970) or the desert forms S. t.
hoggara or S. t. isabellina. Specimens of S. t. turtur in the collection have all
been collected either in their breeding quarters or on migration, so there are
no December specimens for comparison. It would be of great interest to
know where this bird would have wintered. It would seem unlikely that it
was actually on migration in December".
The specimen is a male and had undeveloped gonads and ossified skull. The
fresh dead weight was 105 g and wing length 176 mm.
Cuculus canorus, Grey Cuckoo
A lone bird was sighted in the palm grove by the Research Station on 2
November 1972: this was undoubtedly the individual seen on five subse-
quent occasions, the last being 16 November 1972. Four times the bird was
seen in a large Tournefortia argentea bush at the crest of a small beach. A 'salt
and pepper' moth , Utetheisa lactea, was extremely abundant on this bush and
this insect probably formed a large part of the bird's diet on Aldabra. The
Aldabra Drongo Dicrurus aldabranus was seen to mob the cuckoo severely.
Like the cuckoo recorded for Aldabra by Diamond and Penny (in Benson
& Penny 1971 : 516) this bird appeared as large as any to be seen in England,
which would appear to rule out the possibility of it being the smaller C.
poliocephalus which Benson & Penny consider less likely to occur.
Apus apus, Common Swift
Benson has asked me to include comment on the identity of the specimen
collected by Abbott on 1 December 1892, recorded by Benson (1967: 91) as
A. a. apus. R. K. Brooke (pers. comm. to Benson) re-examined it during a
visit to the Smithsonian Institution in 1968, and reports as follows: — "The
specimen was found to be a juvenile A. a. pekinensis. It agreed completely
with juveniles from Kashmir in lacking the pale patch above the bill which
juveniles of nominate apus have. This difference (in juveniles) is diagnostic
between the two subspecies. The fact that it was a juvenile increases the
chance that the Aldabra bird was lost. No trace of moult in the primaries was
found in this specimen. However, the specimen reported on by Benson &
Penny (1971 : 516), in the British Museum, which has been re-examined by
Benson and P. R. Colston, is still considered to belong to the nominate form.
Telacanthura ussheri, Mottled Spinetail*
Swifts were observed on three occasions at the Old Settlement. They were
about the size of the Common Swift Apus apus, but with distinct white
rumps and square tails. Two birds seen on both 7 and 8 December 1972
(probably the same individuals) were observed to have distinct white vents.
A single individual was seen on 3 January 1973.
I suspected that these birds might be Telacanthura ussheri. The question
was referred by Benson to R. K. Brooke, who has commented that the above
description fits only T. u. stictilaema of eastern Africa or benguellensis of south-
eastern Africa, the former being on geographical grounds the more likely.
He adds that this species is not known to have more than local movements,
and large scale movements are not to be expected.
Merops superciliosus, Madagascar or Blue-cheeked Bee-eater
One was seen, perched in the top of a Casuarina tree by the Research
Station, 15 December 1972. It was not sufficiently closely seen to decide
whether it was a Madagascar Bee-eater M. s. superciliosus or Blue-cheeked
Bee-eater M. s. persicus. Benson & Penny (1971 : 517) give an unequivocal
record of the latter.
15
Eurystomus glaucurus, Broad-billed Roller f
An example was first noted, and collected, at the Old Settlement on 31
October 1972. It proved to be a male, in very fresh plumage. The skull was
ossified, fresh dead weight 119 g, and wing length 210 mm. The testes were
undeveloped, measuring 5 x 3 mm. There were four large insects in the
gizzard; one of both Homoptera and Hymenoptera, and two Scarabaeidae.
The wing length clearly indicates that this specimen belongs to the
Madagascar breeding form, E. g. glaucurus. Benson informs me that two males
and a female which he and A. Williams collected in Madagascar in November
1 972/ January 1973 weighed respectively 136, 166, 149 g. The Aldabra
specimen was evidently emaciated, probably through lack of sufficient
suitable food.
Nine sightings of this species followed the above one, involving at least
six individuals, around the Old Settlement and the Research Station. All were
during November and December 1972, the last being on 19 December. The
kestrel Falco newtoni was seen to mob this species.
All these records are best interpreted as 'fall-outs' on return migration to
Madagascar. Earlier records are discussed by Benson & Penny (1971 : 517),
and for another recent one see Benson (1972).
Hirundo rustka, Swallow
Three appeared on 19 March 1973 and were about the Research Station
for a couple of days, appearing toward evening to hunt over the solar stills.
Previous records of this species on Aldabra are all between 14 March and 10
April (Benson & Penny 1971 : 520).
Motacilla alba, White Wagtail *
D. Bourn reported a single individual at Dune Jean-Louis, South Island,
on 7 March 1973. The bird was observed for a considerable time, and Bourn
made extensive notes and provided me with excellent field sketches. He made
particular note of a grey back and the white on the sides of the head extending
back to meet the grey mantle, and is familiar with the species in Europe.
Anthus trivialis, Tree Pipit * f
This species was first seen near the Research Station on 18 November 1972,
and an individual, no doubt the same one, was collected at the same site two
days later. The specimen is in slightly worn plumage, had a fresh dead weight
of 18 -o g and a wing length of 83 mm. It proved to be a male with gonads
completely undeveloped and skull not quite fully ossified.
Three other pipits were seen, all close to the Research Station in Casuarina
woodland, on 27 November, 17 December 1972 and 13 March 1973. Views
of the first two were poor but they looked like A., trivialis, and the third
was virtually certainly that species.
The specimen confirms that the species does visit Aldabra, and suggests
that the bird seen by Grubb and Hutson (Benson & Penny 1971 : 5 18) at Cinq
Cases in January 1968 was this species.
Oenanthe oenanthe, Wheatear
A grey-backed male was observed about the Research Station from 2 to 4
January 1973. Mrs. Loris ToplirTe informed me of another grey-backed bird
seen inland from the Station on 17 February 1973. Benson & Penny (1971 :
5 19) recorded this species on Aldabra for the first time, and thought it may
regularly occur there in extremely small numbers. My observations tend to
support this contention. For another recent record, see Benson (1972).
Phy/Ioscopus trochilus}, Willow Warbler? *
A bird believed to be a Willow Warbler Phylloscopus trochilus was observed
16
for about five minutes on one of the small islets inside Passe Femme on 10
April 1972. Very good views were obtained as the bird fed on insects from
the foliage of mangrove trees, Rhi^ophora mucronata. It was not collected but
the possibility of it being some other Phylloscopus sp. on geographical grounds
is most unlikely. Furthermore, the Willow Warbler has recently been re-
corded for the Amirante Bank group (Benson 1972).
Phylloscopus sibilatrix, Wood Warbler *f
A single bird was collected at the Old Settlement on 12 December 1972,
whilst feeding on insects from foliage of a Casuarina tree. The fresh dead
weight was 8-6 g, wing length 73 mm. It proved to be a male, in fresh
plumage, with totally undeveloped gonads and a fully ossified skull. Insect
groups represented in the gizzard were, in order of predominance : Hymen-
optera, Dictyoptera, Diptera. This species is an addition to the list by Forbes-
Watson (1972), and Moreau (1972 : 102) gives very few African records from
south of the equator.
Muscicapa striata. Spotted Flycatcher f
A specimen was collected just north of the Research Station on 23 Novem-
ber 1972. It proved to be a male with undeveloped gonads and skull ossi-
fication incomplete. The fresh dead weight was 14-6 g and wing length
88 mm. Contents of the gizzard included insects of the following groups:
Odonata; Coleoptera, Curculionidae; Hymenoptera, Vespoidae; Hemiptera.
Another individual was seen just south of the Old Settlement, 1 December
1972.
Benson & Penny (1 971 : 5 18) give a number of occurrences between 11 and
28 March, which they considered representative of a northward passage.
The above two evidently are referable to a southward one. A specimen
recorded by Benson & Penny was identified as M. s. striata, as is also mine.
Oriolus oriolus, Golden Oriole
One bird, in female plumage, was observed foraging on Casuarina foliage,
19 November 1972, near the Research Station. The only other record of this
species in the Malagasy Region is that of a single bird collected in March on
Aldabra (Benson & Penny 1971 : 521). The earlier record was considered to
be of a lost migrant on northward passage. This November individual
presumably represents a lost southward bound migrant.
Astove Atoll
Eight hours were spent on Astove during 3 April 1973, bird watching in
the coconut palm grove and mixed scrub near the settlement area. Except for
the additional information given below the avifauna was the same as recorded
by Benson (1970).
Phylloscopus trochllus?, Willow Warbler? *
A bird thought to be a Willow Warbler was observed taking insects from
the foliage of a very broad leaved fig-like tree in the centre of the settlement
area. It was in fine plumage and quiet, and very good views were had. This is
the first record of a true land bird migrant for Astove Atoll. Considering also
the records of Phylloscopus spp. already given above, it can be suggested that
warblers are not so very rare on islands in the Malagasy Region, but orni-
thologists are.
Introduced species
Domestic pigeons Columba livia have recently been introduced by the
people resident on the atoll. This is most regrettable as the introduced birds
17
may well preclude the possibility of future colonization by a wild pigeon
species. The present population is estimated at approximately 60 birds. Many
of these can be seen about the settlement (where they are fed), but several
were flushed from the open mixed scrub a considerable distance from human
influence.
Domestic chickens are feral throughout the atoll. Domestic and Muscovy
Ducks Anas platyrhynchos and Cairina moschata are to be found in and about
the settlement buildings.
Farquhar Atoll
During the eleven hours spent on Farquhar on 6 April 1973 a walk from
the settlement area to the far end of North Island and back was made. Habitat
inland was exclusively old cultivated palm grove with shrubs and grasses
beneath. The following records supplement the information in Stoddart &
Poore (1970: 19-21).
Larus fuscus, Lesser Black-backed Gull *
A single individual was seen as soon as the ship anchored off the reef and
observed until the time of departure. The fact that it did not follow the vessel
into the area was confirmed by Mr. Sedgwick, manager and postmaster of
Farquhar, who told me that the bird had been about for over a week. This is
the first record of the species on Farquhar, the second island on which it has
observed in the Malagasy Region, Aldabra being the first (Dawson 1966: 8).
Introduced species
The Barred Ground-dove Geopelia striata was extremely numerous
throughout North Island, parties of five to ten birds being flushed frequently.
Equally numerous was the Madagascar Fody Foudia madagascariensis, many
nests of which were in use.
Other species
Approximately 30 Turnstones Arenaria interpres were noted during the
course of the day.
Acknowledgements
Firstly I thank all my friends and colleagues who were on Aldabra with
me for their enthusuasm in pointing out 'odd' birds. I also thank Antonio
Constance for his assistance in the field; his remarkable eye for wildlife and
accurate descriptions were of great value. For hospitality I am grateful to
Mrs. R. M. Veevers-Carter on Astove, and Mr. Sedgwick on Farquhar. Mr.
B. H. Cogan kindly identified the insects found in gizzards.
The general guidance of Mr. J. F. Peake, Dr. D. W. Snow and Dr. D. R.
Stoddart has been of great benefit. I am also grateful to Mr. C. W. Benson,
Dr. W. R. P. Bourne, Mr. R. K. Brooke, Mr. P. R. Colston and Mr. D.
Goodwin for advice and assistance in the course of the preparation of this
paper. Lastly I must express my appreciation to the Royal Society of London
and those persons on the Society's staff concerned with Aldabra for the
opportunity to visit the atoll and complete my researches on my return to the
United Kingdom.
Summary
Records of migrant and vagrant birds were made on Aldabra during the
period 9 April 1972 to 2 April 1973. Observations on Astove and Farquhar
were made during a day stop at each, en route for the Seychelles from Al-
dabra. Nine species are recorded as new for Aldabra, and one each for Farqu-
har and Astove. Of the species of land birds recorded on Aldabra, all arrived
18
as odd individuals or in very small flocks with the exception of the falcons
which appear to move over Aldabra in a steady migration, mostly in Novem-
ber/December. Large flocks of Crab Plovers Dromas ardeola, observed moving
north from Aldabra, appear to represent a migration to breeding grounds on
the coast of Somalia. There appear to have been some recent introductions
on Astove.
References :
Archer, G. & Godman, E. M. 1937. The birds of British Somali land and the Gulf of Aden 2.
Edinburgh: Oliver & Boyd.
Benson, C. W. 1967. The birds of Aldabra and their status. Atoll Res. Bull. 118: 63-1 11.
— 1970. Land (including shore) birds of Astove Atoll. Atoll Res. Bull. 136: 11 5-120.
— 1972. Some land bird migrants in the western Indian Ocean. Atoll Res. Bull. 162: 10-11.
Benson, C. W. & Penny, M. J. 1971. The land birds of Aldabra. Phil. Trans. Roy. Soc.
Lond. B. 260: 417-527.
Dawson, P. 1966. A survey of the sea birds of the Seychelles Islands. Ool. Rec. 40: 1-11.
Forbes-Watson, A. D. 1972. Preliminary list of the birds of the Malagasy Region. Nairobi:
Nat. Mus. (cyclostyled).
Goodwin, D. 1970. Pigeons and doves of the world. London: Brit. Mus. (Nat. Hist.)
Moreau, R. E. 1972. The Palaearctic- African bird migration systems. London & New York:
Academic Press.
Penny, M. J. 1971. Migrant waders at Aldabra, September 1967-March 1968. Phil. Trans.
Roy. Soc. Lond. B. 260: 549-559.
Stoddart, D. R. 1968. The conservation of Aldabra. Geogr. fourn. 134: 471-485.
— 1 97 1. Scientific studies at Aldabra and neighbouring islands. Phil. Trans. Roy. Soc. Lond.
B. 260: 5-29.
Stoddart, D. R. & Poore, M. E. D. 1970. Geography and ecology of Farquhar Atoll. Atoll
Res. Bull. 136: 7-26.
White, C. M. N. 1965. A revised check list of African non-passerines birds. Lusaka: Government
Printer.
Turtle Dove Streptopelia turtur in South West Africa
by J. M. Winterbottom
Received yth January, 1974
Some years ago, Mr. A. J. Horak, then living on the Okavango River,
caught a strange dove, which he placed in his aviary. Some years later, when
he had moved to Oranjemund, at the mouth of the Orange River, he caught
another. They proved to be female and male respectively and Mr. Horak
succeeded in breeding from them, after the death of the original female, from
the offspring. He has now about eight of these birds, which he identified as
the palearctic Turtle Dove Streptopelia turtur.
I saw these birds on 28 September 1973, and there is no doubt about the
correctness of the identification. The problem is, how did they get to South
West Africa? Vaurie (1965, The birds of the palearctic fauna. N on-pas serif or mes)
gives the winter quarters as "Africa south to Senegal, Gambia, the Sudan
and Abyssinia". The only alternative to their being migrants which have
overshot their mark is that they were escapes from captivity. While this last
is just conceivable for the Oranjemund example, I find it quite incredible that
anybody would be keeping Turtle Doves on the Okavango without Mr.
Horak's knowledge; and we know that other palearctic migrants do over-
shoot their normal wintering range — witness Merlins Falco columbarius in
Natal and European Swallows Hirundo rustica on Marion Island.
[The above note was shown to D. Goodwin, who suggests that some Streptopelia turtur
might go further south than previous observations have suggested. For a record from
Aldabra, see p. 14 above. — Ed.].
19
The significance of records of the Common Sandpiper
breeding in East Africa
by C. If". Benson & M. P. Stuart Irwin
Received 18th November, 1973
As a family the Scolopacidae are virtually confined in their breeding range to
the Palaearctic and Nearctic regions and are usually long-distance migrants.
There are however some major exceptions to this. Members of the Scolo-
pacinae, especially the genus Gallinago, for the most part frequenting swampy
grassland, have managed successfully to colonise the Old and New World
tropics. Meyer de Schauensee (1966) lists five species of the genus as being
endemic to the Neotropical region, where a geographically representative
population of the almost cosmopolitan Common Snipe G. gallinago also
occurs. In Africa the Ethiopian Snipe G. nigripennis, often regarded as
conspecific with gallinago as by Voous (i960), is also widespread on that
continent ; as is the Madagascar G. macrodactyla in the east of that island. In
south-east Asia and Indonesia the Pintail Snipe G. stenura breeds as far
eastwards as the island of Borneo, and the East Indian Woodcock Scolopax
saturata from Sumatra to New Guinea. Two members of the relict tribe
Prosoboniini (classification of Zusi & Jehl 1970: 760-780), one of which is
now extinct, are confined to the Society Islands and the islands of the
Tuamotu group in the tropical Pacific Ocean.
Any other records of scolopacids breeding outside of the Holarctic region
as a whole are therefore of particular zoogeographical interest. Indeed the
only such evidence are records of the Common Sandpiper Tringa hypoleucos in
East Africa. Benson et at. (1970: 14), however, saw no reason for accepting
any African breeding record of this species. Subsequently Cunningham van
Someren (1973) claims that such breeding records are authentic. From the
text of van Someren & van Someren (191 1) it would appear that the first one
is from the shore of Lake Victoria, in Uganda, and that eggs were being
incubated in either August or September. The accompanying photograph of
an incubating bird can surely only be of a Common Sandpiper. It has been
critically inspected by Drs. D. W. Snow and P. J. K. Burton as well as by
Benson himself. There is also a record quoted by Moreau (1966: 124) of
young chicks just fledged seen by Meinertzhagen on the Kajiado River, in
Kenya, in June; and Cunningham van Someren quotes further data from
van Someren (1936: 60-61).
In view of the evidence quoted in the preceding paragraph it is admittedly
going too far to reject these records. Cunningham van Someren indicates that
Benson et at. are following Chapin (1939: 97) and Moreau {loc. cit.) in re-
jecting the evidence. Neither Chapin nor Moreau mentions the data provided
by van Someren (1936: loc. cit.), partly at least a repetition of earlier data. In
fact they did not reject the Lake Victoria and Kajiado records entirely out of
hand. Nevertheless Chapin was sceptical, and Moreau concluded that at most
breeding in East Africa is very rare.
Moreau's conclusion is surely correct, although the Lake Victoria record
seems irrefutable. We do not know of any further evidence of breeding from
anywhere in the Ethiopian region as a whole. The Republic of South Africa
or Rhodesia would seem as likely an area for breeding as any, and in general
is ornithologically the best known part of the Ethiopian region. We therefore
wish particularly to stress that, whatever might have been the situation under
a different climatic regime in the past or might be in the future, in the present
epoch the Common Sandpiper cannot be regarded as in any sense a regular
member of the Ethiopian breeding avifauna. It would appear to us that the
case of the Common Sandpiper is in part analogous to that of the White
Stork Ciconia ciconia and House Martin Delichon urbica, recorded as breeding
occasionally in South Africa, as discussed by Moreau (1966: 123), only in
this instance it is the sole member of the Tringinae to breed anywhere within
the tropics.
Doubtless Cunningham van Someren is correct in stating that Common
Sandpipers are present throughout the year in East Africa. We can corrobo-
rate from our own experience in Rhodesia and Zambia, merely as one further
example, that this is also the case further south. But a similar situation exists
for some other scolopacine waders, e.g. the Greenshank Tringa nebularia, and
is not evidence of local breeding. Thus Dowsett (in Benson et al. 1970: 14)
counted more than 500 Greenshanks in a stretch of 300 miles of the Luangwa
River walked in July and August 1966 and 1967. Actually Dowsett (pers.
comm.) finds that the Greenshank is the commonest of the scolopacines which
overwinter in Zambia, the Common Sandpiper the least so, and so the least
likely to breed ; this presumably applying in other parts of Africa too.
Finally, we cannot but remark, that despite the amount of ornithological
investigation in East Africa in recent decades, no further evidence in support
of these earlier observations has come to light.
We are grateful to R. K. Brooke and R. J. Dowsett for their comments
on a draft of this note.
References :
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. Stuart. 1970. Notes on the
birds of Zambia: Part V. Arnoldia {Rhodesia) 4(40): 59 pp.
Chapin, J. P. 1939. The birds of the Belgian Congo. Part II. Bull. Amer. Mus. Nat. Hist.
75: 632 pp.
Cunningham van Someren, G. R. 1973. Tringa hypoleucos Linnaeus breeding in East Africa.
Bull. Brit. Orn. CI. 93: 39-40.
Meyer de Schauensee, R. 1966. The species of birds of South America. Livingston: Wvnewood,
Pa.
Moreau, R. E. 1966. The bird faunas of Africa and its islands. London: Academic Press.
van Someren, V. G. L. 1936. The birds of Kenya and Uganda. Journ. E. Afr. <& Uganda
Nat. Hist. Soc. 13 (1, 2): 29-72.
van Someren, R. A. L. & van Someren, V. G. L. 191 1. Studies ofbirdlife in Uganda. London :
John Bale & Danielson.
Voous, K. H. i960. The Atlas of European birds. London: Nelson.
Zusi, R. L. & Jehl, J. R. 1970. The systematic relationship of Aechmorhynchus , Prosobonia
and Phegornis (Charadriiformes ; Charadrii). Auk 87: 760-780.
The predators of the Jackass Penguin Spheniscus demersus
by J. Cooper
Received 19 th September, 19J3
INTRODUCTION
Temperate zone penguins appear to have different predators and may ex-
perience a lower level of predation than sub- Antarctic and Antarctic species
(Stonehouse 1967). Important predators in high latitudes are the Leopard
Seal Hydrurga lepontonyx, Great Skua Catharacta maccormicki, and Giant Petrel
Macronectes giganteus (Stonehouse i960, 1967; Swales 1965; Young 1963,
1970). Virtually nothing is known of the predators or the level of predation
of temperate species.
Casual unquantified observations were made of predation on the temperate
Jackass Penguin Spheniscus demersus, and on its eggs and chicks during the
period February 1971 to December 1972 on DassenIsland(33°2 5'S, i8°o6'E)
off the west coast of South Africa. Predation of the Jackass Penguin is
considered in three sections ; predation of eggs, chicks and j uveniles and adults .
PREDATION OF EGGS
The only observed predator of eggs was the Dominican Gull Larus
dominicanus. The gull was never observed to take eggs from incubating
penguins as does the Great Skua (Young 1970). However the possibility
exists that gulls may work in pairs to entice penguins from their nests
(Headman, Dassen Island, pers. comm.). Gulls were only seen to take eggs
from deserted surface nests and were therefore acting more as scavengers
than as predators. Human disturbance to breeding penguins permits gulls to
obtain more eggs than they would under undisturbed conditions. Deserted
eggs in burrows were left untouched, since they were invisible to gulls
patrolling overhead. The increase in the penguin's surface nesting habit due
to guano removal and paving of the breeding areas on some islands has
increased the gulls' chances of obtaining eggs. Berry et al. (in press) found
that gull numbers increased on Halifax Island, South West Africa, during
guano collection and that many eggs were taken by gulls as the penguins
deserted because of human disturbance. Guano removal has apparently
caused an increase in surface nesting on Halifax Island.
While skuas and Giant Petrels occur in the seas around Dassen they are
not seen ashore, and therefore do not take eggs (or chicks) as they do in high
latitudes.
Man has been an important predator of Jackass Penguins. From 1917 to
193 1 an average of 460,000 eggs was officially collected annually on Dassen
Island (Cott 1953). Collecting was halted in 1968 and poaching is now at a
low level, but still causes some disturbance.
PREDATION OF CHICKS
Penguin chicks are predated by Dominican Gulls, Sacred Ibis Threskiornis
aethiopicus and possibly by feral domestic cats Felis lybica. Gulls took small
unattended chicks (up to one week old) from surface nests. Human disturb-
ance causing temporary desertion increases chick loss since small chicks are
normally brooded continuously. Larger chicks were only attacked if very
weak. The Sacred Ibis was only seen to attack weak unattended chicks,
mostly comatose. This bird mainly scavenges corpses, and is only a casual
predator.
Feral cats occur on Dassen Island feeding mainly on the feral rabbit
Oryctolagus cuniculus. It is possible that they sometimes take penguin chicks
though this was never proved by stomach content examination. Small chicks
often disappeared from deserted burrows inaccessible to gulls and could have
been taken by cats. One penguin chick (ca. 1,500 g) was found showing signs
of having been partially eaten by a cat.
Chicks have been collected for food and for fish bait in the past, but man
is no longer a predator. The extinction of the penguin colony on Robben
Island in Table Bay (South Africa) was probably partially due to the collection
of chicks in large numbers.
PREDATION OF JUVENILES AND ADULTS
The Cape Fur Seal Arctocephalus pusillus was the only predator seen to take
fledged penguins. A single animal, thought to be a female by its size, was
often seen feeding on penguins in House Bay, Dassen Island. A total of 57
penguin corpses was washed up, but many more must have been carried
away by prevailing winds and currents. On one day a total of 25 was washed
up. The seal's technique was invariably to decapitate the bird with one bite
and then to disembowel the carcass by shaking, removing the belly skin and
eating the viscera and stomach contents. The breast muscle and the rest of
the corpse were left untouched. The Cape Fur Seal is not thought to be a
common predator of the Jackass Penguin; Rand (1959) mentions only one
bull seal which was feeding on penguins off Dyer Island near Cape Agulhas
in 1937. A similar case occurred at Halifax Island in 1957. Bourne & Dixon
(1973) quote an observation made at sea near Dassen Island where "a seal
was seen to catch a penguin, shake it about and toss it in the air". This was
most probably a Cape Fur Seal. The habit seems to be confined to individuals
which only appear interested in the stomach contents of penguins. The seal
was not always present and it is considered unlikely that penguins formed its
sole diet as no drastic decline in penguin numbers was noticed at the landing
stages in House Bay.
On other islands seals and penguins apparently live together in harmony.
I have observed seals close inshore at Dyer Island and Possession Island,
South West Africa, but they were not seen to catch penguins. Predation of
penguins by Fur Seals is not considered regular by Stonehouse (1967).
However, the New Zealand Fur Seal A. forsteri takes Rockhopper Penguins
Eudyptes crestatus in quantity off Campbell Island (Bailey & Sorensen 1962),
and apparently on Macquarie Island as well (Warham 1963). Boswall (1972)
quotes observations by Ian Strange of the South American Sea Lion Otaria
byronia killing the Magellanic Penguin Spheniscus magellanicus in the Falkland
Islands. Sea lions do not occur in South African waters. The Leopard Seal is
very rare in South African waters (P. Best pers. comm.) and thus cannot be
an important predator of the resident Jackass Penguin.
An old record of a Bryde's Whale Balaenoptera edeni (—brydei), with 15
Jackass Penguins in its gut (Olsen 191 3), is considered to be unusual (P. Best
pers. comm.), as this animal has a narrow gullet and feeds on small fish and
euphausids. Bird remains were absent from 119 Bryde's Whale stomachs
examined from South African waters (Best 1967).
Killer Whales Orcinus orca have been recorded taking penguins in high
latitudes and do occur in South African waters. Penguins have not been
foundin ten stomachs examined by P. Best although one contained an albatrosse.
The Killer Whale is probably only a minor predator of Jackass Penguins.
Giant Petrels have been seen scavenging on Jackass Penguins killed by
Cape Fur Seals but have not been seen to attack live birds; at Gough Island
this petrel kills freshly moulted Rockhopper Penguins as they enter the
water (Swales 1965).
Likely, but unproven predators of penguins are sharks. Penguins were
often seen on Dassen Island with missing feet or truncated flippers. These
injuries could have been caused by seals or sharks. Hagen (1952) mentions
the Blue Shark Glyphis glaucus as a possible predator of Rockhopper Penguins
at Gough Island.
Historically man has been an important predator of juvenile and adult
penguins. In the seventeenth and eighteenth centuries many birds were col-
lected for food on Robben Island. They have been used as fish bait, and are
occasionally drowned in fishing nets. At present man is no more than a casual
predator of Jackass Penguins.
SUMMARY
The Jackass Penguin is preyed upon by several animals, but none of these
appears to be an important predator. Predation of eggs by Dominican Gulls
is related to the level of human disturbance and the increase in surface nesting.
The Dominican Gull and Sacred Ibis take small unattended or weak chicks.
Feral domestic cats may also take chicks, but proof is lacking. Juvenile and
adult birds are preyed on by individual Cape Fur Seals but this is a rare
occurrence. The Cape Fur Seal was only observed to eat viscera and stomach
contents. Predation by sharks is an unknown factor, and Killer Whales,
Leopard Seals, Skuas and Giant Petrels are not thought to be important
predators in South African waters. Man has been an important predator of
penguins in the past, taking eggs, chicks and fledged birds, causing the
extinction of at least one island breeding colony. Predation by man is no
longer important.
AC KNOWLEDGEMENTS
During my period of research on Dassen Island I was employed by the
South African National Foundation for the Conservation of Coastal Birds
under a grant from the South African Nature Foundation. I am grateful for
facilities on the island supplied by the Department of Government Guano
Islands. The paper was prepared while I was a member of the Percy Fitz-
Patrick Institute of African Ornithology.
References :
Bailey, A. M. & Sorensen, J. H. 1962. Subantarctic Campbell Island. Denver Mus. Nat.
Hist. 10: 1-305.
Berry, H. H., Seely, M. K. & Fryer, R. E. in press. The status of the Jackass Penguin
Spheniscus demersus on Halifax Island off South West Africa. Madocqua.
Best, P. B. 1967. Distribution and feeding habits of baleen whales off the Cape Province.
Div. Sea Fisheries Invest. Report 57: 1-44.
Boswall, J. 1972. The South American Sea Lion as a predator on penguins. Bull. Brit. Orn.
CI. 92: 129-132.
Bourne, W. R. P. & Dixon, T. J. 1973. Observations of seabirds, 1967-1969. Sea Swallow
22: 29-60.
Cott, H. B. 1953. The exploitation of wild birds for their eggs. Ibis 95 : 409-449.
Hagen, Y. 1952. Birds of Tristan da Cunha. Norweg. Sci. Exped. to Tristan da Cunha 20.
Olsen, O. 191 3. On the external characteristics and biology of Bryde's Whale (Balaenop-
tera brydei), a new rorqual from the coast of South Africa. Proc. Zoo I. Soc. Lond. 34:
i-75-
Rand, R. W. 1959. The Cape Fur Seal, Arctocephalus pusillus, distribution, abundance and
feeding habits off the south western coast of the Cape Province. Div. Sea Fisheries
Invest. Report 34: 1-75.
Stonehouse, B. i960. The King Penguin, Aptenodytes patagonica, of South Georgia. 1.
Breeding behaviour and development. Falkland Islands Depend. Sci. Report 23: 1-8 1.
— 1967. The general biology and the thermal balance of penguins. Advances in Ecol.
Research 4: 1 31-196.
Swales, M. K. 1965. The seabirds of Gough Island. Ibis 107: 17-42.
Warham, J. 1963. The Rockhopper Penguin, Eudyptes chrysocome, at Macquarie Island. Auk
80: 229-256.
Young, E. C. 1963. Feeding habits of the South Polar Skua, Catharacta maccormicki. Ibis 105 :
301-318.
— 1970. The techniques of a skua-penguin study. Antarctic Ecology 1 : 568-584.
The Boran Cisticola in Ethiopia
by J. S. Ash
Received jth Jul) , 1973
Following Benson's (1946) account of unidentified Cisticolas in southern
Ethiopia, North & McChesney (1964) collected and recorded what were
apparently similar birds in Kenya. The latter authors dubbed them "Boran
Cisticolas".
Although Boran Cisticolas are morphologically inseparable from the
24
Rattling Cisticola C. chiniana (Hall & Moreau 1970) their behaviour, song,
and habitat preferences are dissimilar ; clearly, two species are involved.
At the risk of possibly confusing the problem further, it seems worthwhile
recording some observations on a Cisticola which seems to be fairly wide-
spread in southern Ethiopia. I have used the same name Boran Cisticola for
these birds, although I cannot be certain that they are the same as those
described by the above authors. This seems likely, for they are inseparable in
the field from chiniana on plumage characters, but differ in other respects.
Their most noticeable feature is the loud song, quite unlike the rattling song
of chiniana, and to my ear strongly reminiscent of the palearctic Chaffinch
Fringilla coelebs.
Boran Cisticolas have been found most commonly in the Kebre Mengist
and Neghelli areas, in Borana, part of Sidamo (Benson recorded them at
Neghelli). In April 1971 they were fairly common between these two places
at 75 km by road (5°3i'N, ^°z^'K) east of Kebre Mengist, and here a bird
was incubating four eggs on 1 7 April in a ball-shaped nest ca 1 5 cm above the
ground in roadside weeds; they were again common at 40 km to the east
(o5°i4 / N, 39°57'E) and at 54 km to the south (a$°oo'N, 39°29'E) of Neghelli.
In April 1973 they were common at various points between these localities
and at Neghelli itself. On 23 April 1971 by the Genale River (o5°32'N,
39°42'E), north of Neghelli, a bird left a nest at 0700 which contained a warm,
and probably new-laid egg; there was still one egg at dusk, but two at 0800
the following morning. This nest was similar to the other, and was ca 25 cm
above the ground in low herbaceous growth. Benson recorded a nest with
four eggs on 19 June. Since April 1971, further birds with the same charac-
ters of plumage and song have been observed, and these records suggest
that the species is widespread in southern Ethiopia: Gibe gorge (o8°o8'N,
37°29'E), Kaffa Province: fairly common on 30 May and 19/20 June 1971;
Dire Dawa, Harar Province: one singing at 10 km to south (o9°34'N,
4i°5 2'E) on 5 September 1971 ; Harar, Harar Province: single birds at 26 km
to east (o9°i4 / N, 42°i8'E) and 46 km to east (o^^'N, 42°28'E); Metahara,
Shoa Province: single birds at 62 km and 66 km to west (o8°4i'N, 39°3o'E);
Langano (7°34'N, 38°5i'E), Shoa Province: one on 3 October 1971, and 6+
singing on 13 February 1972 and 6/7 May 1972, and subsequently over a
fairly wide area in 1972 and 1973, singing in March, May and June; Arba
Minch (6°o2'N, 37°36'E), one on 14/15 April 1972; Yavello, at 15 km to the
north-east (5°oo'N, 38°i2 / E), one on 2 June 1972; Ghinir (7°8'N, 4o°43'E),
common on 9 April 1973.
Whereas there are only a few records of the Boran Cisticola in the Rift
Valley, most of my records of chiniana are confined to that region at many
localities from Bahadu (io°o5'N, 4o°37'E) in the north to Lake Chamo
(5°48'N, 37°3i'E) in the south. Outside the Rift Valley chiniana has only been
found at Yavello and Neghelli. Nests with eggs have been found in April,
July and September. There are also records in the south at Bulcha (6°27'N,
38°n'E), near Lake Margherita (=Abbaya), but these birds have not been
heard singing, so it is uncertain if they are chiniana or the Boran, although I
have specimens.
Boran Cisticolas tend to be found in thicker and lusher cover than chiniana,
and they have only been found within sight or sound of each other in three
areas. At Arba Minch in April 1972, one bird was singing in a thick cover of
bushes on top of the escarpment, whilst on the slopes, 50-100 m below,
chiniana was common. At one Langano site, Boran Cisticolas occur in the
thick cover at the base of the escarpment on the lake-shore, separated by
2 5
only about a kilometre from the nearest chiniana in the more open acacia bush,
but at another site they occur together. At Yavello, a singing Boran was in
an area where chiniana was common. The Boran has been observed at an
altitude range of 879-1573 m, and chiniana at 600-2000 m.
To summarise my observations on the range of the two species: C. chiniana
extends throughout most of the Rift Valley, only occurring beyond this
area locally in Borana. Urban & Brown (1971) state that it occurs north to
Eritrea, where it is rare. The Boran overlaps the range of chiniana broadly
in the Rift Valley, except north of 8°3o'N, with extensions to the north-east,
east and west. Further observations, especially in the south-west, may alter
this picture. However, on extensive journeys further east in Haud, Ogaden,
southern Sidamo and Bale, there has been no sign of either species.
Morphological differences between the two species are described by Erard
in the immediately following paper.
Acknowledgements : I am grateful to C. Erard, K. D. Smith and Dr. C. K.
Wallace, for commenting on this paper. Part of these observations were
made during the course of research supported by the Office of Naval Re-
search Task Order No. N00014-67-A-0 3 99-0009 and Bureau of Medicine and
Surgery Work Unit MR 041.09.01 - 0014D6HJ. The work was undertaken
under the auspices of Dr. George E. Watson of the Smithsonian Institution
and Drs. J. R. Schmidt and C. K. Wallace of U.S. Naval Medical Research
Unit No. 3, Research Detachment, Addis Ababa, Ethiopia.
References :
Benson, C. W. 1946. Notes on the birds of southern Abyssinia. Ibis 88: 180-205.
Hall, B. P. & Moreau, R. E. 1970. An atlas of speciation in Ajrican passerine birds. London:
Brit. Mus. Nat. Hist.
North. M. E. W. & McChesney, D. S. 1964. More voices of African birds. Cornell University:
Laboratory of Ornithology.
Urban, E. K. & Brown, L. H. 1971. A checklist of the birds of Ethiopia. Addis Ababa: Haile
Sellassie I University.
The problem of the Boran Cisticok
by C. Erard
Received 20th August, 1973
INTRODUCTION
The existence in Ethiopia of two sibling species of cisticolas was first brought
to light by Benson (1946: 199-203). This author collected in Sidamo Cisticola
chiniana (Smith) and another species morphologically similar but different in
voice and habitat. Later, North & McChesney (1964) collected and recorded
them also, in Kenya, at Marsabit (2°2o'N, 37°5o'E) and Nanyuki (o°oi'N,
37°o5'E), and gave the vernacular name "Boran Cisticola" to the second
species.
In 1968, during one of the expeditions to Ethiopia organised by the
Laboratoire de Zoologie (Mammiferes et Oiseaux) du Museum National
d'Histoire Naturelle de Paris, J. Prevost and the author in their turn found
two cisticolas in Sidamo which, although morphologically similar, differed
in voice, behaviour and biotope. Still not knowing well the cisticolas of
Ethiopia and not being conversant at the time with the problem posed by
Benson's observations, they thought that there must be obvious differences
which would be apparent when skins were compared. Also, detailed notes
26
on the voice of each specimen collected were not made systematically. This
was an error. The material collected in 1968 remained unidentified.
In 1 97 1, during another expedition, Prevost and the author, accompanied
by N. Follet, took up the problem again. For each specimen collected a
record was made as to its type of voice, while at the same time a series of
specimens of the two species was collected which brought to light morpho-
logical differences. In the discussion which follows, this sibling of the Rattling
Cisticola C. chiniana will be referred to as the Boran Cisticola, which can be
regarded as of established usage.
VOICE
This is the best criterion for distinguishing the two species, and it is
doubtless this which plays the most important part in their isolation from
one another. Lynes (1930: 247) has described well the voice of chiniana. The
alarm cry can be rendered as "ptit" or "tchit", sharp and repeated, sometimes
"tititi" as in a tit Parus sp. The song consists of two or three notes, detached
and a little drawn out (most frequently two notes, somewhat grating,
reminiscent of a Whitethroat Sylvia communis in tone), followed by a short
trill. In the field, it was recorded as "piu-piu-trrtrrtrr" (the most frequent),
"tu-tu-tititi", "tu-tu-titititi", "prr-prr-vouivouivouivoui". The same song as
in Ethiopia was heard in December 1971 in Kenya and Tanzania.
The alarm cry of the Boran Cisticola is a nasal "tsee-tsee", sometimes
repeated in a series of as many as ten such notes. It is usually the male which
makes this call, facing the intruder, on a conspicuous perch, a special posture
being adopted. He stands erect, making himself as tall as possible, the wings
being beaten spasmodically and vigorously, and brought to the vertical
at each beat, while at the same time balance is kept by violent vertical jerks
of the tail. This has the effect of dispersing a family group. A playback on
the magnetophone of these calls causes the bird to emit more in a louder
tone; if this is continued, it becomes more and more excited, sings and attacks
the magnetophone. This would suggest that the calls do not only serve to
warn the partner and their offspring but also have a combative value in
territory maintenance. In fact, when a male penetrates the territory of another,
he is evicted with cries similar to his own. We have also heard, rarely, from
excited birds a dry "tec-tec", sometimes followed by a rapid "tsuisuisui".
The song of the Boran Cisticola recalls, to a European ear (see also Ash
1974), certain stanzas of the Chaffinch Fringilla coelebs, especially at the end of
winter when they are not yet complete. This song is a rapid, very loud
"rolling" which one could even qualify as explosive (see also Benson loc.
cit.). The phonetic transcriptions of the song in nature are remarkably
constant: "til-lilulululu" or "tchit-tilulululu" or "tip-tilululu", the indi-
vidual variation being relatively slight and depending especially on the
length of the stanza.
Figure 1 shows the sonagrams of the songs of the two species. It appears
that the birds recorded by North in Kenya are indeed Boran, not Rattling
Cisticolas. A comparison, kindly carried out and commented on by Dr. C.
Chappuis, shows that: —
(1) In true chiniana the song is composed of a regular initial series of vibrant
notes, long and of a complex tonal structure. This is followed by one or two
series of short but identical notes of a rapid rhythm.
(2) In the Boran Cisticola according to North and in Ethiopia, the song is
composed of an irregular initial series of notes modulated in frequency, short
and of a simple tonal structure. This is followed by a series of notes of varying
frequency and higher intensity.
27
&l ,
I !
2-
Hh MM *t|. WMt * II "I
B
I . I
h % If fc
4-
Ili4i
D
05
i r
10 1-5
Seconds
20
2-5
Figure i. Wide-band sonagrams showing the following songs: — A, B and C, different
forms of the song of the Rattling Cisticola; D. Boran Cisticola (by North).
KHz
8-1
4-
KHz
2-
j
$ & m if
<
I MM
1 I 1
! i i *****
—
KHz
8n
KHz
8n
6-
I
1^
05
Mm
10 1-5
Seconds
20
2-5
Figure i {continued). The same showing: — E, song of Boran Cisticola (by North); F and
G, song, H, alarm cry, of Boran Cisticola (by author and Follet, at Kebre Mengist).
Dr. Chappuis adds that, comparing North's recordings and ours, in general
the songs are identical : the same structure of the initial notes, much the same
number of notes in the second part of the phrase, the same form of the notes.
Furthermore, the initial notes of the song of Boran Cisticolas in Ethiopia are
absolutely identical with those in Kenya. Thus one may conclude finally that
these birds constitute a single species, quite distinct from the Rattling
Cisticola.
In spite of the fact that our material was not very adequate, we may re-
count our experience in replaying the song of the one species to the other
near Neghelli. When we played to chiniana the alarm cry followed by the song
of the Boran Cisticola, it replied with cries and some song, but the reactions
did not attain the intensity, not to say violence, provoked when its own song
was played to the Boran. On the other hand, when the cries and song of
chiniana were played to the Boran, a positive response of cries and song were
evoked. The difference in the intensity of the replies is perhaps explained by a
difference in the biological cycles : the fact that chiniana was at the finish of the
postnuptial moult and was singing relatively little, whereas the Boran,
although it had certainly also finished breeding, had only just started to
moult and was still singing quite well. This confirms the suspicion of Hall &
Moreau (1970: 167) that chiniana has an earlier breeding season than the
Boran. Our observations also showed clearly that the two species, when they
occur alongside one another, are mutually exclusive territorially. It would
seem, although our data are too fragmentary to confirm it, that chiniana and
the Boran can have contiguous territories but not actually overlapping.
HABITAT
Benson (1946: 201), in Sidamo, has stressed that chiniana inhabits typically
"the open 'thorn-acacia' country at 3000-4500 ft.", whereas the Boran lives
in "the drier highlands at 5000 ft. and over ... in open country where there
is low bush growth". In general we confirm this difference in habitats. We
found chiniana only in dry thorny savannas, always in areas where there are
relatively low and scattered shrubs, interspersed with low, thick bushes,
thinly scattered. At Arero, however, we observed and collected it in culti-
vated hollows, partly fallow, where there were only some poorly developed
bushes on the edge of a streamlet.
On the other hand, the Boran inhabits dense tree savannas of a dry type,
though also fairly humid as for example at Kebre-Mengist in Sidamo. Thus
it is common in glades and on the outskirts of light forest below 1800 m, in
open evergreen shrubby growth, in degraded vegetation on the edges of
roads, in man-made clearings reminiscent in appearance of the Mediterranean
garrigue (as in the Gibe valley in Kaffa, where it is also found in quite open
places). In general, it frequents fairly dense but broken shrubby formations.
Around Neghelli and towards Arero and Yavello, and doubtless beyond,
the two species come into contact, and in certain places live side by side in
the same habitat. However, closer observation shows that chiniana prefers the
poorer, more open areas, in which it is much more abundant than the Boran,
which dominates on the other hand when the height and density of the woody
vegetation increases. Thus for example near Neghelli and Yavello they both
occupy thornbush, but the Boran prefers the more luxuriant spots (dense
shrubby or bushy growth with a more herbaceous floor), whereas chiniana
prefers the more open spots in which the acacias are reduced to dispersed
clumps and grass exists only in strands around such clumps. The Boran sings
especially at the tops of tall acacias (song post between 3 and 5 m, or even
30
higher), whereas chiniana sings low down in bushes or on the tops of grass-
stems (song-post between o- 5 and 2- 5 m, rarely higher). It would also seem
that then; is a difference in the feeding zones : the Boran living between 1 and
5 m and foraging between 1 and 3 m, with chiniana keeping preferably to
below 2 m and foraging essentially in grass and low branches of bushes
between ground-level and 1 m.
As to altitude, we do not think that there is any clear-cut difference between
the two species. Chiniana was found between 920 and 1620 m, the Boran
between 1200 and 1800 m (the altitudes were taken with the aid of an
altimeter previously adjusted in the Geophysical Laboratory, Haile Sellassie
I University, Addis Ababa).
MORPHOLOGY
The examination of specimens of the two species duly identified by voice
has permitted the recognition of a certain number of differences in colour,
form and measurements. Thus the series brought back in 1971 show the
following differences in the Boran Cisticola from the Rattling Cisticola: —
(a) The back is more uniform, less streaked. The basic colour of each
feather is brown, clearly lighter and with a pronounced russet tone. The
streaks of the mantle, although practically non-existant in worn plumage, are
narrower and above all browner, less black.
(b) The crown is browner, less of a warm chestnut, almost unstreaked,
while in chiniana it is obscured by dark streaks.
(c) The underparts are whiter, less buff.
(d) The tenth (outermost) primary is shorter (see measurements below).
(e) The wing-coverts (especially the greater and median) and the alula are
bordered with whitish or very light buff and contrast with the reddish buff
edges to the outer webs of the remiges. In chiniana the wing-coverts and the
remiges are fringed with rufous or reddish fawn uniformly dark.
(f) The rectrices are narrower. On the under surface, the extremity beyond
the subterminal black spot is practically of the same colour as the basal part
of the feather (see also Benson 1946 : 201), whereas in chiniana this apical zone
is white or washed with fawn, and contrasts clearly with the colour of the
base of the feather. The subterminal black spot in the Boran is in fact no more
than blackish, somewhat diluted as remarked by Benson (Joe. cit.) y and is
concentrated on the inner web, being absent or almost so on the outer. In
chiniana this spot is more definitely black, and is equally well defined on both
webs, so that it may fairly be termed a bar. This black marking is also less
extensive in the Boran than in chiniana (less than 5 mm in extent, as against
6-8 mm). In the Boran, too, it is barely discernible on the upper surface,
whereas in chiniana it is still clear. Furthermore, in the Boran black is entirely
absent on the central pair of rectrices, while in chiniana a rudimentary vestige
is still visible.
(g) The bill is shorter, more thickset at the base, resembling somewhat in
profile, in miniature, that of Cisticola nataiensis (Smith).
The above differences being evident in autumn specimens (Boran starting
to moult, chiniana finishing), an attempt was made to find them in specimens
in fresh dress. A comparison of our material with that in the British Museum
(Natural History), wherein is the series collected by Benson (most of which
are Boran, as their collector thought) has confirmed the validity of the above
31
findings. In fresh plumage the Boran Cisticola has however the wing specu-
lum (edges of the greater and median coverts) a little more buff than in worn
plumage, but nevertheless this speculum contrasts with the edges of the
remiges. The underparts are decidedly whiter than in chiniana in the same
stage of plumage, in which they are much washed with buff, with white only
in the centre of the abdomen and of the throat. Furthermore, chiniana has on
the sides of the chest a dark greyish wash which tends to make it appear
darker on the underparts. The pale edges of the mantle feathers in the Boran
in fresh dress have a greyish rather than fawn tone, and are narrower than in
chiniana, which explains why, with wear the back in the Boran appears the
more uniform of the two species.
Until the present, these two cisticolas were placed together in a single
species as Cisticola chiniana bodessa Mearns, cf. White (1962: 663). Thanks to
the great kindness of Prof. S. Dillon Ripley and Dr. R. L. Zusi, we have
examined the type of this form, described under the name C. subruficapilla
bodessa by Mearns (191 3: 2-3). It shows all the characters of the Boran
Cisticola, the scientific name of which must accordingly be Cisticola bodessa
Mearns. The populations of chiniana of Ethiopia and northern Kenya must
be called C chiniana fricki Mearns, of which we have also examined the type
(cf. Mearns 191 3: 3-4), and which corresponds well with specimens which
we have collected after having identified them from the song as Rattling
Cisticolas.
The birds of Kaffa (40*6*, J 9) differ from the other Boran Cisticolas, and seem
sufficiently distinct to bear their own particular name:
Cisticola bodessa kaffensis subsp. no v.
Description: This form differs from the nominate by its clearly darker
colour. The crown and the mantle are almost concolorous. The top of the
head, of a less warm tone than in true bodessa, is scarcely more rufous than
the mantle, which is a duller, darker, brown, lacking any rufous ; in addition
the mantle is practically uniform even in fresh dress, at the most only with
very ill-defined streaking. The primaries are bordered with a more pro-
nounced, darker, reddish, while the margins of the lesser and median wing-
coverts are browner, less greyish. The thighs are likewise a darker buff. The
underparts appear darker, being slightly more buffy and, more especially,
more washed with a darker brownish grey on the flanks and sides of the
chest. Also, the extremity of each rectrix is lighter, more washed with buff,
in contrast to the rest of the feather. The bill seems to be longer.
Distribution: So far only known from the valley of Gibe in the province of
Kaffa.
Type: $ adult: between Walkite and Abalti, ca. 8°i2'N, 37°4o'E, 7 June
1968. Collector's no. 1438. In the Museum National d'Histoire Naturelle de
Paris. Wing 67, tail 63-5, bill from skull 14 mm; weight 20 g.
The material collected from Marsabit (30*0* m the British Museum) is not
separable from nominate bodessa.
To return to our comparison between chiniana and bodessa, in measure-
ments the two species are very similar. The following figures show such
differences as do exist (each set of figures shows respectively the mean, the
range in brackets and the number of specimens examined): —
32
Wing
SS 66-5(60-71)
43
99 53>55
2
$$ 66-0(64-68)
4
Tail Bill from
skull
Weight
millimetres
grammes
C. b. bodessa
57-1 (52-63) 13-5 (12
5-I4-5)
19-5 (16-
-«)
4i 39
27
48, 50 11, 13
12, 13
2 2
2
C £. kaffensis
58-6 (52-5-63-5)14-0 (13
5-i5)
20 -o (19-
-")
4 4
4
45 13
14
1 1
1
C. chiniana fricki
57-3 (52-62) 14-6 (13-
-16)
20-6 (18-
-23)
37 37
21
50-8 (50-53) 13-0 (12-
-14)
14-8(12-
-20)
5 6
5
S? 52
I
o*o* 67-0(60-70-5)
39
99 55-2(51-5-58)
6
In order to take better account of structural differences between the two
species, we have shown on histograms the frequency of length of wing and
bill of each (Fig. 2). The similarity of size is reflected by length of wing.
Chiniana is only slightly larger than bodessa, but on the other hand there is a
more definite difference in length of bill.
In the course of collecting, we were struck by the difference in the relative
length of the tenth (outermost) primary. To express the validity of this, we
have measured the difference in length between the extremity of the tenth
primary and that of the tip of the wing (tip of the longest - usually the seventh
- primary). Figure 3 shows that at an equal wing-length bodessa has in general
a shorter tenth primary, although there is a considerable zone of overlap.
This is reduced if one compares bill-length to the percentage of difference
between the tip of the wing (longest, seventh primary) and the tip of the
tenth primary in relation to the wing-length. This percentage is another way
of expressing the relative length of the tenth primary to the full wing-length.
These two structural differences in the wing and the bill, combined with the
colour differences already described, permit the identification of single,
isolated specimens.
MOULT
Lynes (1930: 270-271), not knowing that there were the two species,
qualified as "irregular" the mode of moult, stating that in Arussi, Sidamo
and the Rift Valley, from Lake Zwai to Stephanie (that is, in the zone where
both species exist) there were two modes, seasonal and perennial, although
elsewhere the seasonal mode was "dominant if not exclusive" (see also
Friedmann 1937: 200). According to the specimens which we collected and
those in the British Museum, it appears that bodessa has a complete moult
between the end of October and the beginning of January, and chiniana from
September to mid-December. This is general, and affects all of the two popu-
lations. A bodessa in moult of the primaries was obtained on 22 June at
Marsabit by North, and a chiniana starting to moult on the River Gato (near
Gardula) on 3 1 August. Indications of "spring" moult have been noticed in
specimens of bodessa as follows : one in moult of the primaries at Neghelli in
33
No. specimens
15-
10
S
60 61 62 63 64 65 66 67 68 69 70 71 Wing (mm)
Figure z. Above: Wing-lengths of Cisticola bodes sa (vertical lines) and of C. chinianafricki
(horizontal lines) {$$ only). Below: Bill-lengths of C. bodessa (continuous line) and of C. c.
fricki (broken line) (o*6* only).
No. specimens
15-
10-
5-
I 1
12
2.5 1 13 ' 13.5 1 14 '14.5 1 15 ' 15-5 1 16 '
Bill (mm)
34
2/
26 H
25
24-
23-
22-
21-
20
19-
18-
17-
Wing tip— 10 primary (mm)
60 61 62 63 64 65 66 67 68 69 70 71 Wing (mm)
Figure 3. Above: Relation between wing-length and difference between tip of wing and
tip of tenth (outermost) primary {$<$ only). Below: Relation between bill-length, and per-
centage of difference between tip of wing and tip of outermost primary to wing- length
(o*6* only). In both graphs C. bodessa is shown by circles, C. chinianajricki by triangles. Solid
symbols: identification by song. Hollow symbols: identification from specimens only.
Underlined symbols: C. b. kaffensis.
Bill (mm)
16-
15-
14
13
12-1
n 1 1 1 1 1 1 1 1 1 1 1 r
28 29 30 31 32 33 34 35 36 37 38 39 40
Wing tip — 10 primary /
Wing
35
Sidamo on 15 May; two showing signs of moult of the rectrices, secondaries
and body plumage in KafTa on 7 and 19 June; likewise in chiniana, two
specimens showing moult of the median rectrices at Lake Margherita in the
Rift Valley on 28 April and 2 May. This "spring" moult seems to affect only
a very small number of individuals, contrary to the "autumn" moult, which
appears to be the true moult of the populations as a whole. Further investi-
gation in the field is necessary to establish clearly the cycle of moult of the
two species ; all the more so because the cycles of reproduction and moult
must certainly vary locally in accordance with the incidence of the rains.
DISTRIBUTION
Ash (1974) has outlined the distribution of the two species in Ethiopia
according to his personal observations which it is not necessary to discuss
here. We give below a summary of our own by provinces, adding to it
information obtained from the material in the British Museum. We will not
consider the data obtained by North in Kenya which also concerns the Boran
Cisticola.
Eritrea: A £ from Nefasit (1 5°i9'N, 39°o4'E), 8 June 195 1, despite its bad
state of plumage, seems to be a bodessa. Smith (1955 : 75, under C. chiniana
bodessa) states "Scarce and local ... on eastern escarpment between 4000-
6000 ft". It would be desirable to examine a series of specimens of this
apparently isolated Eritrean population.
Harrar: A <$ bodessa was collected by Pease on 15 December 1900 at
Hojojo, west of Mieso (ca. 9°io'N, 4o°4o'E). Ogilvie-Grant & Reid (1901:
654), writing of this specimen, have already remarked that it differs from
others in having the streaking of the back almost obsolete.
Shoa: There are specimens of bodessa collected along the Awash River on
19 December 1900 by Pease and at "Manniballa" on 14 July 1902 by Degen.
This latter locality must be the same as the "Mallabella" (ca. 9°io'N, 39°4o'E)
of Pease if reference is made to the itineraries of the two expeditions (Ogilvie-
Grant & Reid 1 90 1 : 609 and Ogilvie-Grant 1904: 254). The Boran Cisticola
was also well represented on 1 April and 10 October 1971 in the region of
Welenchiti (8°4o'N, 3?°25'E) and on 1 April at Sodere (8°2o'N, ^o'E).
Chiniana was heard singing (it was not at all rare) in many localities in the
Rift Valley during May, June, July, September, October and December in
1968-71, especially between Modjo (8°39'N, 39°o5'E) and Shashemene
(7°i2'N, 38°37'E). It was also heard near Nazareth (formerly Adama,
8°3i'N, 39°i2'E) and near the crater lake of Debre Zeit (8°48'N, 38°5 5'E).
We have examined specimens collected in February from Jawaha (ca. io°N,
4o°E), a locality investigated by Blundell and Lovat, see Ogilvie-Grant
(1900: 118), in April from Dembi (visited by Antinori and probably in the
Rift mid-way between Addis Ababa and Lake Zwai), in January from Lake
Zwai (ca. 8°oo'N, 38°5o'E) and in September from Kambata (ca. 7°3o'N,
38°2o'E), to the south-west of Lake Shalla (see Ogilvie-Grant 1913 : 555 and
pi. 12).
Arussi: We have examined specimens of chiniana collected by Zaphiro in
January/February at Djila, Dalota and Bourka (localities to the east of Lake
Zwai, between this lake and the high valley of the River Cathar; see Ogilvie-
Grant 191 3 : 554).
Sidamo: It is in this province that the two species seem most abundant.
Chiniana was found around Lake Awasa (7°N, 38°25'E) in May/June 1968,
then in large numbers in a radius of 15-20 km around Neghelli (5°2o'N,
39°3 5'E) in May 1968 and October/November 1971, thence very commonly
36
in November/December 1971 to Arero (4°48'N, 38°5o'E), onwards almost
throughout to Yavello (4°5 7'N, 38°o8'E), and for at least 50 km to the west
and 70 km to the south thereof (and see also Benson 1946: 200).
Bodessa was abundant in May 1968 and October/November 1971 at Kebre
Mengist (formerly Adola, 5°54'N, 38°5 9'E) and 50 km east of Neghelli
(ca. 5°io'N, 4o°o5'E) towards Filtu. In fact its density was highest between
Wadera(5°4o'N, 3 9°2o'E) and 15 km north of Neghelli (ca. 5°2 3 'N, 39°3 5'E),
in this region indeed to the exclusion of chiniana (see also Ash's records). In
November/December 1971, the Boran Cisticola was equally well represented
around Arero, and in the vicinity of Yavello from 10 km to the north thereof
to 75 km to the south (that is, where Benson op. cit. found it himself).
Gemu-Gofa: We have examined specimens of bodessa dated June and
August from Konso, between Lakes Shamo and Stephanie, for June from
Wurke (ca. 6°3o'N, 37°3o'E, see Ogilvie-Grant 1913: 554), for May from
Bodessa (5°o5'N, 37°32'E, see Friedmann 1930: 7-8), and for January from
Muji, to the north of Lake Rudolf (this locality is not shown on any map;
specimen obtained by Haverstadt). Above the escarpment dominating the
western shore of Lake Margherita, at several points, between Soddu (6°49'N,
37°4i'E) and Arba Minch (6°o2'N, 37°36'E), we heard the song in May 1968.
In this same region chiniana was common at the bottom of the escarpment,
on the edge of the lake. The type offricki, belonging to this latter species,
came from "White Abaya Lake" ( — Lake Shamo), and was collected on 20
March 191 2. We have also examined a series of specimens of chiniana from
Konso, in the valley of the River Gato, collected by Zaphiro in August 1910.
Kaffa: In June 1968 and November 1971 we found bodessa well represented
in the Gibe valley between Walkite (8°i5'N, 37VE) and Abalti (8°io'N,
37°3 5'E), and above the escarpment or a further 6 km to the south-west of
Abalti. It was apparently in this same area that Ash found the species in May
and June 1971. In June 1968 singing was heard in clearings near Agaro
(7°5o'N, 36°38'E), but not again in November 1971. In the meantime the
habitat had been considerably modified by the construction of an all-season
road, accompanied by cultivation and human activity in general.
AC KNOWLEDGEMENTS
I must express my gratitude to my travelling companions Dr. J. Prevost
and N. Follet, without whom this study would have been impossible. I wish
also to thank Dr. J. S. Ash, who helped me with his knowledge of Ethiopian
birds; Dr. D. W. Snow and I. C. J. Galbraith, who gave me every facility
while I was working in the British Museum (Natural History) at Tring; and
Prof. S. Dillon Ripley and Dr. R. L. Zusi, who were kind enough to lend me
the type specimens described by Mearns. Special mention must be made of
C. W. Benson, who has commented on this paper and translated it into
English; and of my friend Dr. C. Chappuis, who made the sonagrams and
from whom I have profited by his knowledge of birds in general and of
cisticolas in particular. Mrs. J. Hall-Craggs also most kindly assisted in the
final preparation of the sonagrams before they were sent to the printer;
likewise J. W. Rodford in that of Figures 2 and 3.
SUMMARY
Based on differences in voice and to some extent in ecology, it was sus-
pected more than 30 years ago that there were two species of cisticola in
southern Ethiopia. In the meantime, both have been known as Cisticola
chiniana bodessa. The present investigation has confirmed these differences,
37
and has also shown that there are small differences in colour of plumage and
in structure. The two species may be known as the Rattling Cisticola C.
chiniana, widespread in southern and eastern Africa, and extending into the
Shoa province of Ethiopia; and the Boran Cisticola C. bodessa, known from
Nanyuki and Marsabit, northern Kenya, and widespread in Ethiopia,
occurring even in Eritrea. A new subspecies of the latter, C. b. kaffensis, is
described from the Gibe valley, Kaffa. The main factor keeping the two
species apart is voice. In some places (as in Sidamo) they occur alongside
one another, but are territorially mutually exclusive.
References (excluding those in the paper by J. S. Ash above) :
Ash, J. S. 1974. The Boran Cisticola in Ethiopia. Bull. Brit. Orn. CI. 94: 24-26.
Friedmann, H. 193 0-1937. Birds collected by the Childs Frick expedition to Ethiopia and
Kenya Colony. Smiths. Inst. U.S. Nat. Mus. Bull. 153. Part 1. Non-passeres: 516 pp.
Part 2. Passeres: 506 pp.
Lynes, H. 1930. Review of the genus Cisticola. Ibis 12(6), suppl.: 673 pp.
Mearns, E. A. 191 3. Description of six new African birds. Smith. Misc. CoWns 61(11): 1-5.
Ogilvie- Grant, W. R. 1900. On the birds collected during an expedition through Somaliland
and southern Abyssinia to the Blue Nile. Ibis 7(6): 11 5-178, 304-337.
— 1904. On the birds collected during a recent expedition through Somaliland and
Abyssinia to lake Tsana. Ibis 8(4): 250-280.
— 1913- On a collection of birds from southern Abyssinia presented to the British Museum
by Mr. W. N. McMillan. Part I. Passeres. Ibis 10(1): 550-641.
Ogilvie-Grant, W. R. & Reid, R. J. 1901. On the birds collected during an expedition
through Somaliland and southern Abyssinia to lake Zwai. Ibis 8(1): 607-699.
Smith, K. D. 1955. Recent records from Eritrea. Ibis 97: 65-80.
White, C. M. N. 1962. A check-list of the Ethiopian Muscicapidae (Sylviinae). Part II.
Occ. Pap. Nat. Mus. S. Rhod. 26 B: 653-694.
Cape Grass Owl in Ethiopia
by D. A. Turner
Received )oth October, 197)
The Cape Grass Owl Tyto capensis ranges through the greater part of southern
Africa northwards to western and central Kenya, where it is a rather rare
and local species frequenting damp open grassy areas. Mackworth-Praed &
Grant (1970) also indicate the existence of an isolated population on Manen-
guba Mt., western Cameroun. Urban & Brown (1971) do not list this species
for Ethiopia, and Leslie Brown informs me that even the Barn Owl Tyto alba,
widely distributed over most of Africa south of the Sahara, is unaccountably
rare in Ethiopia.
On 9 May 1973 I observed an owl asleep on a telegraph pole in the middle
of a vast open grassy area near Tefki, some 50 km south-west of Addis
Ababa, Ethiopia. The bird was observed for some 1 5 minutes at a distance
of approximately 75 yards, with the aid of x 8 binoculars. It was clearly a
Tyto owl, the facial discs being very distinct, as were its orangey-buff under
parts lightly spotted with dark brown or black. The upper parts and wings
were uniform dark brown. The area at this time was extremely dry, although
it was very apparent that it became a vast swampy area during the rainy
season. When I left, the owl was still asleep, but had disappeared when I
returned to the area later in the afternoon.
A recent examination of skins in the National Museum, Nairobi, of all
medium sized owls leaves me with no doubt that the bird I saw was a Cape
38
Grass Owl. This would appear to be the first record of this species in Ethiopia
and a considerable northward extension of its previously known range.
References :
Mackworth-Praed, C. W. & Grant, C. H. B. 1970. African handbook of birds, Series I. Vol. 1.
London: Longmans.
Urban, E. K. & Brown, L. H. 1971. A checklist oj the birds of Ethiopia. Addis Ababa: Haile
Sellassie I Univ. Press.
Locustella naevia in Ethiopia
by J. S. Ash & George E. Watson
Received pth October, 1973
Whilst one of us (J.S.A.) was routinely trapping birds and mammals at Koka
(8°27'N, 39°o6'E) at 1445 m in Shoa Province, Ethiopia, in connection with
research on viral infections, a Grasshopper Warbler Locustella naevia was
netted at 11 30 on 24th February 1973. It weighed 10 • 2 g, wing length 62 mm,
and was judged on the basis of colour to belong to the eastern race straminea.
Subspecific identification was confirmed by John Farrand, Jr., at the Smith-
sonian Institution, where it is deposited (USNM 552733).
Previously only one other Ethiopian record of L. n. straminea has been
reported, namely from Abroberifaghe along the River Hawash in the Danakil
in February 1947 (Guichard 1948). Until the more recent records from Sene-
gal (Moreau 1972) this has remained the only known occurrence south of the
Sahara. However, we have been unable to trace this previous Ethiopian
specimen. The record is included without date or authority in Meinertzhagen
(1954), Mackworth-Praed & Grant (i960), White (i960) and in the recent
Ethiopian checklist (Urban & Brown 1971). Moreau (1961), however, cites
Meinertzhagen (Joe. cit.) as authority for the record but he later (1972) gives
Thesiger & Meynell (1935). The species, however, is not mentioned by
Thesiger & Meynell. There is no voucher specimen in either the British
Museum or Meinertzhagen collections (D. W. Snow, in litt., 17 May 1973)*.
C. W. Benson (in lift., 21 July 1973), who checked independently the British
Museum collections, also failed to find the specimen. He searched the regis-
ter and found that three of the four species covered by Guichard's (pp. cit.)
note were duly recorded, but not the Locustella. One might assume that the
specimen in question never reached the museum if it were not for the state-
ment at the end of Guichard's note, "All the above-mentioned specimens are
now in the British Museum". Alternatively, the specimen may have been lost
prior to registration. This supposition is supported by the fact that only
eight out of the nine Sarothrura ayresi sent to the museum were registered.
Guichard was a careful and reliable observer, and notwithstanding the fact
that his specimen of L. naevia cannot be traced now we are inclined to accept
it. The species is not included in the catalogue of the collection made during
the Italian occupation (Italian Game Department, unpublished, photostat
copy of disintegrating original in J.S.A.'s possession).
Moreau (1972), in discussing the wintering range of races of L. naevia,
follows others in identifying the earlier problematical Ethiopian bird as L. n.
straminea, mentioning that both this and another eastern race, L. n. mongolica,
*A. D. Forbes-Watson (pers. comm.) asked Thesiger in Kenya on 22 June 1973 about the
problematic record. Thesiger had no recollection of having collected the species and stated
that if the record was not cited in the 1935 paper, nor was there a specimen in the British
Museum, it never in fact existed.
39
winter regularly in India (Vaurie 1959). Moreau's review of records in the
western Sahara and Senegal does not include mention of the race involved,
although one at least was the European race L. n. naevia (Roux 1959), to
which they are presumably all referable. Field (1973) has reported recent
observations of Grasshopper Warblers from further south in Sierra Leone,
West Africa.
The present bird was in dry. Acacia jBalaniteshush ca. 1/2 mile from the lake
shore. Other Palaearctic migrants, many of-which must have over-wintered,
caught in the same habitat at the same time, included: 11 Hoopoes Upupa
epops, three Scops Owls Otus s. scops, two Wrynecks Jynx torquilla, five Pied
Wheatears Oenanthe pleschanka, two Stonechats Saxicola torquata armenica, 24
Redstarts Phoenicurus p. phoenicurus and samamisicus, three Bluethroats
Luscinia svecica magna, three White-throated Robins Irania gutturalis, 13
Nightingales Luscinia megarhynchos, 12 Lesser Whitethroats Sylvia curruca,
three Whitethroats S. communis, seven Blackcaps S. atricapilla, four Barred
Warblers S. nisoria, four Olivaceous Warblers Hippolais pallida elaeica, 24
Reed Warblers Acrocephalus scirpaceus, one Willow Warbler Phylloscopus
trochilus, 29 ChiffchafTs P. collybita, eight Masked Shrikes Lanius nubicus and
four Red-tailed Shrikes Lanius collurio isabellinus.
During 1970-73, 9639 birds have been netted on 70 days at the Koka site
(6 in February, 29 in March, 19 in April and 16 in December), and at least
2624 (27%) of these were migrants, but no other Grasshopper Warblers
were seen. Similarly, much netting has been done in other sites in southern
Ethiopia, including two in the Danakil, all of which have provided many
migrants, but not L. naevia. Unless the species is confined largely to small
areas of specialised habitat, which have not been worked yet, we suspect it
cannot be more than a rare and casual winter visitor to East Africa south of
the Sahara.
The opinions and assertions in this scientific report are those of the authors
and do not necessarily reflect the official views of the Navy Department or of
the naval service at large. The research was supported in part by the Bureau
of Medicine and Surgery and the Office of Naval Research under Contract
No. Nooo 1 4-67-A-03 99-0009 and Bureau of Medicine and Surgery Work
Unit MR041.09.01-0014DGHJ. We are grateful to C. W. Benson, J. Farrand,
Jr., the late K. D. Smith and D. W. Snow for their help.
References :
Field, G. D. 1973. Subalpine and Grasshopper Warblers in Sierra Leone. Bull. Brit. Orn. CI.
93 : 101-103.
Guichard K. M. 1948. Notes on Sarotbura {sic) ayresi and three birds new to Abyssinia.
Bull.' Brit. Orn. CI. 68: 102-104.
Mackworth-Praed, C. W. & Grant, C. H. B. i960. African handbook of birds. Series I, Vol. 2.
London: Longmans.
Mcincrtzhagen, R. 1954. Birds of Arabia. Edinburgh & London: Oliver & Boyd.
Morcau, R. E. 1961. Problems of Mediterranean-Saharan migration. Ibis 103a: 373-427,
580-623.
— 1972. The Palaearctic- African bird migration systems. London & New York: Academic
Press.
Roux, F. 1959. Captures de migrateurs palcarctiques dans la basse vallee du Senegal. Bull.
M//s. Nat. Hist. Nat. Paris ze(^i): 334-340.
Thesiger, W. & Meynell, M. 1935. On a collection of birds from Danakil, Abyssinia. Ibis
77: 774-807.
Urban, E. K. & Brown, L. II. 1971. A checklist of the birds oj Ethiopia. Addis Ababa. Haile
Scl lassie I Univ. Press.
Yauric, C. 1959. The birds of the Palearctic Fauna: Order Passeriformes. London: Witherby.
White, C, \I. N. i960. A check list of the Ethiopian Muscicapidae (Sylviinae). Part I. Occ.
Pap, Nat. Mus. S. Rhod. 3(24!}): 399-430.
40
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/ 1
Bulletin of the
British Ornithologists' Club
c
Edited by
C. W. BENSON
Volume 94 No. 2
June 1974
Committee
J. H. Elgood {Chairman)
P. Hogg {Vice-Chair man)
Sir Hugh Elliott, Bt., o.b.e. {Editor)
R. E. F. Peal {Hon. Secretary)
M. St. J. Sugg {Hon. Treasurer)
J. H. R. Boswall
Dr. C. J. O. Harrison
C. J. Mead
Lt.-Col. J. R. Neighbour
Bulletin of the /^
BRITISH ORNITHOLOGISTS' CLUB
Vol. 94 No. 2 Published: 20 June, 1974
The six hundred and eighty-seventh meeting of the Club was held at the
Cafe Royal, 68 Regent Street, London, W.i, on Tuesday 19th March 1974
at 7 p.m.
Chairman: Sir Hugh Elliott, Bt., o.b.e. ; present 22 members and 1 1 guests.
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Annual General Meeting
The eighty-second Annual General Meeting of the British Ornithologists'
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Regent Street, London, W.i. Eight members were present.
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The Hon. Secretary presented the Report of the Committee for 1973
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There being no nominations additional to those of the Committee, the
following were declared elected : —
Chairman: Mr. J. H. Elgood {vice Sir Hugh Elliott, who retired on com-
pletion of his term of office).
Vice-Chairman: Mr. P. Hogg {vice Mr. J. H. Elgood).
Editor: Sir Hugh Elliott, Bt., o.b.e. {vice Mr. C. W. Benson, o.b.e., who
retired on completion of his term of office).
Hon. Secretary: Mr. R. E. F. Peal (re-elected).
Hon. Treasurer: Mr. M. St. J. Sugg {vice Mr. P. Tate, who retired on com-
pletion of his term of office).
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Hugh Elliott had done for the Club as Chairman. In particular, he had
41
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said that the meeting was well aware of its debt to Mr. Benson and it was
hoped that he would be willing to resume office as Editor in 1975.
It was proposed by Mr. D. T. Holyoak and seconded by the Hon. Secretary
that the meeting be adjourned until 6 p.m. on Tuesday 17th September 1974
at the Cafe Royal, 68 Regent Street, London, W.i for consideration of the
Accounts for 1973, and this was carried unanimously.
The meeting was adjourned at 6.30 p.m.
A re-examination of material of the extinct marabou
stork, Leptoptilos falconeri: with descriptions
of some new species
by C. J. O. Harrison
Received 1st February, 1974
Two marabou storks have been named as existing in pre-pleistocene periods.
Argala (= Leptoptilos) arvenensis was listed by Milne-Edwards (1871) as a
species of the French Miocene, but it is a nomen nudum. No material is
known and it is possible that proposed specimens may have been referred to
other species.
In the same work Milne-Edwards (1867-71), in a footnote to page 450
(incorrectly stated to be 449 in Brodkorb 1963), proposed the name Argala
falconeri for another species. Brodkorb (1963) lists this also as a nomen nu-
dum and regards Davies (1880) as the first valid description. He appears to
have overlooked that on page 449 Milne-Edwards refers to four specimens
from the Siwalik Hills of India, in the collection of the British Museum
(Natural History) and that the name is then proposed with reference to these.
They are listed as the distal and proximal ends of a tarsometatarsus, and two
distal ends of tibiotarsi, measurements are given, and the greater size in
comparison with Recent species is indicated as a valid character for specific
separation. Leptoptilos falconeri (Milne-Edwards) 1868 would therefore
appear to be the original description.
Davies (1880) refers to five specimens, four presented to the British
Museum (Natural History) by Col. Sir P. T. Cautley in 1 842, and one pre-
sented by Charles Falconer in 1868, under this name. Lydekker (1884) refers
a further three specimens to this species, two from Cautley and one from
Falconer, and in describing the largest distal end of the tibiotarsus says it
"may be considered as the type'*. In a later catalogue (1891) he lists all
specimens without special reference to this one; but Brodkorb (1963) lists
it as the designated lectotype. It is an unfortunate selection in that the
specimen, apart from its larger size, shows no useful characters for separation
from Recent species.
This material referred to L. falconeri has been re-examined and I am of the
opinion that it comprises the remains of three different bird species. Five
42
specimens, including the lectotype, appear referable to a very large marabou
stork under the existing name. Although there is no indication that these
were originally associated, the similarities of relative size and structure
appear to justify referring them to a single species.
The lectotype tibiotarsus shows no obvious characters, apart from a
much larger size, which would separate it from the corresponding bone of the
larger Asiatic species of the Holocene, L. dubius. The other, paratypical
specimens show distinct differences in some points of structure which
would justify specific separation, and in which the Pliocene bird is more
different from the three Recent species than they are from each other.
In the comparison of Pliocene and Holocene material this similarity in
one bone and differences in others may have particular relevance. Although
it has been claimed that some Recent species existed in the Pliocene, Brod-
korb (1971) contends that the Pliocene birds were specifically distinct and
that in general Recent species evolved during the Pleistocene. If the Pliocene
records are based on single, or few, skeletal elements then a similar situation
might occur to that apparently shown by the present species, where a
particular bone might give no indication of difference, although differences
existed in other parts of the skeleton, and where it might be assumed that a
Recent species had been in existence at a much earlier period when a mor-
phologically distinct, although possibly ancestral, form might be involved.
The three remaining specimens consist of two distal ends of tibiotarsi,
one very incomplete, and the extreme proximal end of a tarsometatarsus with
most of the hypotarsus. One tibiotarsal fragment is virtually indistinguish-
able in size and shape from those of the Recent JL. dubius. The other more
extensively damaged fragment (fig. 4) is that of another stork, possibly
related to Ciconia species but not definitely referable to known Recent or
extinct species, and apparently a new form. It is from a large bird, only
slightly smaller than L. dubius. The tarsometatarsal fragment appears to be
that of a marabou (fig. 3), in some respects most similar to L. crumeniferus
of Africa. Since, as in the larger species, it is this and not the tibiotarsus that
shows characters which might justify specific separation I regard this as the
holotype and refer the first of the two smaller tibiotarsi to it.
Rich (1972) has assigned a distal end of a tarsometatarsus and a tibiotarsus
from the Upper Miocene Beglia Formation of Tunisia to Leptoptilos cf.
falconeri. The species was a small one, intermediate in size between L.
crumeniferus and L. javanicus. The tarsometatarsus shows a trochlea for the 3rd
digit shorter and broader in proportion than that of L. fa/coneri, having a
distally short trochlea for the 2nd digit with a posteriorly and proximally
displaced outer rim (fig. 1, lower right; 2, top). The tibiotarsus has a larger
and disto-proximally longer posterior articular surface in the intercondylar
region. There seems no reason to assign this bird to L. falconeri other than
on its age. It appears to represent a discrete form and I have therefore
treated it as a new species, L. richae.
The three Recent species oiLeptoptilos show small interspecific osteological
differences. The fossil forms discussed here show, even on the fragmentary
material available, more distinct differences which are usually greater than
the more recent interspecific variations. It therefore seems best to treat all
these earlier forms as separate units distinguished by name.
The relative similarity to each other of the three Recent species within the
genus as a whole suggests that they are probably derived at a later period
from a common ancestral stock, probably diverging during the period of
active speciation in Africa and Asia during the Pleistocene. In addition we
43
Figure i. Anterior view of distal end of tarsometatarsus of -
top left, jL. dubius; top right, L. javanicus; bottom left, jL.
falconeri; bottom right, L. richae (the last, after Rich 1972,
is on scale approx. half as large again as others).
Figure 2. Lateral, internal view
of distal end of tarsometatarsus
of- top, jL. richae; middle, JL.
falconeri; bottom, L. dubius (the
first, after Rich 1972, is on scale
approx. half as large again as
others).
Figure 3. L. shvalicensis, holc-
type, proximal end of tarso-
metatarsus. Internal and proxi-
mal views.
Figure 4. Cryptociconia indica,
holotypc, distal end of tibic-
tarsus. Anterior and internal
44
have two Pliocene and one Miocene species. There is some evidence that on
the fossil forms the anterior condylar projections of the distal ends of
tibiotarsi are slightly longer, relative to the width of the ends, than on
Recent species; but apart from this the scanty material available does not
appear to indicate any general evolutionary trends, unless one interprets the
variation in the distal end of the tarsometatarsus (fig. i) as evidence of a
long-term tendency for an increase in the distal projection of the trochlea
for the 2nd digit. It is possible that these earlier forms are close to the
evolutionary stem of the genus, but there is no definite evidence to indicate
whether any should be regarded as ancestral to the Recent species or whether
they should be regarded as minor divergences from a main evolutionary
trend. There is certainly no good evidence to justify the creation of palaeo-
species.
SPECIES DESCRIPTIONS
Leptoti/osfa/coneri (Milne-Edwards) 1868
Emended diagnosis. Very large. On tarsometatarsus, trochlea for third digit
large, extending distally well beyond other trochleae. Trochlea for second
digit posteriorly displaced. Distal end of humerus with shaft less inflated
anconally; groove between attachment of anterior articular ligament and
internal condyle broad and deep; entepicondylar prominence and attachment
of anterior articular ligament large. Distal end of femur with rotular groove
deep and narrow and external edge of popliteal hollow well-defined. First
phalanx of second digit of manus with metacarpal facet deep relative to
width.
Material. Lectotype, designated by Lydekker (1884), a distal end of a right
tibiotarsus, B.M. (N.H.) no. 39735 (erroneously quoted by Brodkorb 1963
as 39753). Paratypes, a distal end of a left tarsometatarsus, B.M. (N.H.) no.
39736; a distal end of a left femur, B.M. (N.H.) no. 39737; and a proximal
portion of the first phalanx of the second digit, B.M. (N.H.) no. 39738; all
the above specimens including the lectotype being presented by Col. Sir
P. T. Cautley in 1842. Also a distal end of a left humerus, B.M. (N.H.) no.
48435, presented by Charles Falconer in 1868.
Occurrence. Siwalik Series, Lower Pliocene; Siwalik Hills, Uttar Pradesh,
India.
Description. The lectotype is the distal end of a right tibiotarsus with a short
portion of the shaft (fig. 1, lower left; 2, middle). It has undergone some
pressure, the internal condyle showing a slight inward deflection and some
damage to its outer edges. The shaft has been subject to crushing and
deformation, and anterior projections proximal to the condyle are super-
ficially eroded. The characters it shows do not differ distinctly from those of
the Recent species, being perhaps most similar to those of L. javanicus.
The condyles are stout and rounded anteriorly and posteriorly, and concave
on the anterior outer surfaces. The internal ligamental prominence is large.
There is a large, rounded and relatively deep intercondylar hollow distal to
the supratendinal groove.
The distal end of a left tarsometatarsus is broken off at the metatarsal facet.
It is in good condition with the posterior tips of the outer trochleae and the
internal posterior edge of the trochlea for the 2nd digit showing some wear
and erosion. In general it is very similar to the tarsometatarsus of Recent
species but the trochlea for the 3rd digit is larger, extending further distally
and more prominent anteriorly. The anterior edge of the external rim of the
trochlea for the 2nd digit is slightly more prominent laterally.
45
The distal end of the left femur is broken off just proximal to the popliteal
area. The fibular groove and posterior intercondylar fossa are worn and the
posterior external condyle edge slightly eroded. Other surfaces show small
areas of damage. The rotular groove is deep and well-defined, narrower in
comparison to those of Recent species, with a prominent and anteriorly
tapering posterior external condyle. The popliteal cavity is deep and appears
to have a deep and well-defined inner margin, the external portion of the
shaft forming a definite ridge bordering it. This does not appear to be the
result of distortion.
The proximal part of the first phalanx of the second digit consists of a
little over a third of the bone, with the flange mostly broken away. The
proximal end is stout, with the metacarpal facet deep relative to its width.
The distal end of a left humerus is again very similar to those of Recent
species. It appears to lack any suggestion of proximal anconal inflation of the
shaft however. The attachment surface for the anterior articular ligament
forms a blunt and palmarly prominent ridge in these birds and in the present
species is particularly prominent; while the entepicondyle appears to show
greater lateral projection.
Measurements. L.ectotype. Greatest length 52-8; width and depth of shaft
(damaged) at level of proximal opening of supratendinal bridge 20-4 x 16-8;
width at distal end 28-0; width of posterior condylar area 22-9; anterior/
posterior thickness of intercondylar groove 22-6; anterior length of external
condyle 24-8; anterior/posterior width of internal condyle 34-0, of external
condyle 34-8; width of tendinal bridge 8-i mm. Tarsometatarsus. Greatest
length 42-6; proximal width and thickness 23. 1 x i*o; length from mid
proximal end of shaft to trochlea for 2nd digit 39-5, to trochlea for 4th digit
39-9; anterior length of trochlea for 3rd digit 25-4, posterior length iyo,
dorsal width 13-4, greatest depth 17-5; same measurements for 2nd digit
19-8, ii-i, io-i, 17-8; for 4th digit 21-6, 12-0, 10*4, 18-7; greatest distal
width 36-0; dorsal distal foramen to external intertrochlear notch 8-2 mm.
Femur. Greatest length 41-5 ; greatest width 46-8; anterior/posterior width of
external condyle 41-2; distal width of rotular groove 16-1, proximal width
13-8; anterior/posterior thickness of external side 35-3, of internal side
29-0 mm. Phalanx. Greatest length 29-2; width of shaft at distal end 12*1;
thickness of external side 5-8; width of anterior surface at metacarpal facet
15-7; depth of facet 13-1; posterior/external to anterior/internal edges of
facet 15-0 mm. Humerus. Greatest length of fragment 66-9; width and
greatest thickness of proximal end of shaft 35-7 X 20-7; greatest distal width
57-7; thickness at external condyle, 29-0, at internal condyle 19-8; attachment
of anterior articular ligament to ridge of internal tricipital groove 27*5 mm.
l^eptoptilos siwalicensis , sp. nov.
Etymology. The specific name is derived from that of the range of hills in
which the specimens occurred.
Diagnosis. Moderately large. Proximal end of tarsometatarsus with internal
side stout. Hypotarsus only slightly posteriorly elevated. Anterior ends of
calcaneal ridges abrupt. Slanting muscle attachment surface at proximal
posterior edge of external side large.
Material. Holotype a proximal end of a left tarsometatarsus, B.M. (N.H.) no.
39741, presented by Col. Sir P. T. Cautley in 1842 (fig. 3). Paratype a distal
end of a right tibiotarsus, B.M. (N.H.) no. 39734, from the same source.
Occurrence. Siwalik Series, Lower Pliocene; Siwalik Hills, Uttar Pradesh,
India.
46
Description. The holotype is the extreme proximal end of a tarsometatarsus
with the greater part of the hypotarsus present but most of the anterior
surface missing except in the region of the cotyla (fig. 3). It has a general
similarity to those of Recent species, but the internal side at the proximal end
is stouter, anterio-posteriorly thicker, where it borders the hypotarsus. The
base of the hypotarsus appears smaller, the whole structure projecting less
posteriorly. Perhaps partly as a result of this the calcaneal ridges arise more
abruptly and the area of posterior surface immediately around them is
flatter. The attachment surfaces on the posterior and anterior edges of the
proximal end of the external side are large.
The tarsometatarsus is slightly smaller than that of the Recent L. dubius.
The distal end of a tibiotarsus is very similar to that of the latter species, very
slightly larger in the condylar region and with a stouter shaft. Compared
with those of Recent species the anterior portions of the condyles, particu-
larly that of the internal condyle, are longer and narrower relative to the
width of the distal head.
Measurements. Tarsometatarsus. Greatest length of fragment 34-5; greatest
proximal width 28-6; width at cotyla lips 24-3 ; posterior projection of hypo-
tarsus, to rim of internal calcaneal ridge 19-5; to external calcaneal ridge
16-7; posterior external muscle scar 9-6 x 6-8 mm. Tibiotarsus. Greatest
length of specimen 61-4; anterior length of internal condyle 18-9; of external
condyle 19-0; anterior/posterior thickness at internal condyle 30-6, at inter-
condylar groove, 19-4, at external condyle 31-6; greatest distal width 24-4;
posterior length of external condyle 16-6; width and thickness of shaft
c. 35 mm proximal to condyles 15-3 x 13*8 mm.
Leptopti/os richae, sp. nov.
'Etymology. The species is named after Mrs. Pat Vickers Rich who described
the specimens.
Diagnosis (based on Rich 1972). Tarsometatarsus with trochlea for the third
digit relatively short and anteriorly broad. Trochlea for the second digit
does not extend as far distally and has the outer rim much reduced and
posteriorly and proximally displaced. Internal intertrochlear notch wide.
Distal end of tibiotarsus has posterior articular surface of intercondylar
region distoproximally long. Internal condyle projects anteriorly beyond
external condyle. In external view posterior edge of shaft straight.
Material. Holotype a distal end of a right tarsometatarsus, no. T-3604,
Colorado Tunisian Collection, Service Geologique, Tunis. Paratype a distal
end of a right tibiotarsus, no. T-1396, in the same collection.
Occurrence. Lower Faunal Level (localities 17 and 20); Beglia formation.
Upper Miocene; Bled ed Duoarah, Tunisia.
Description (based on Rich 1972). The holotype is a distal end of a right
tarsometatarsus, broken across diagonally a little below the metatarsal facet,
and lacking the trochlea for the fourth digit and the lateral side proximal to it
(fig. 1, lower right). The trochlea for the third digit is narrow posteriorly
with a proximal taper; but anteriorly it is broad in relation to its length. The
internal intertrochlear notch is wide, although this is partly modified
anteriorly by the projecting flange of the trochlea for the third digit. The
trochlea for the second digit is laterally and posteriorly displaced with a
distinct external angle at the proximal end. Its concave inner side extends
distally for about three-quarters the length of the other trochlea, but its
external facet is reduced, and proximally and posteriorly displaced (figs.
1, lower right; and 2, top). The distal end, viewed anteriorly, shows a strong
47
proximo-external slant, and in lateral view a prominent posteriorly-directed
flange is apparent.
The distal end of a tibiotarsus shows the typical rounded anterior and
posterior condylar surfaces. The posterior, intertrochlear articulation surface
is distoproximally longer than that of other species, and the posterior
condylar flanges larger. In distal view the posterior and anterior edges of the
internal condyle appear more anterior relative to the external side. In lateral
view the posterior of the shaft, proximal to the condyles, is straight. Anteri-
orly the tendinal canal is not internally deflected where it enters the proximal
opening of the supratendinal bridge. The distal opening of the bridge is
more medially placed than in Recent species and the internal ligamental
prominence is level with the distal edge of the supratendinal bridge and not
distal to it.
Measurements (after Rich 1972). Holotype. Distal width of trochlea II 6-4;
of trochlea III 9-8; internal depth of trochlea II 13-0, of trochlea III 16-0;
lateral depth of trochlea II 13-5, of trochlea III 16-4; posterior length of
trochlea II 11*9, of trochlea III 15-7; anterior length of trochlea II 17*6, of
trochlea III 23-2, width across trochleae II and III 26-0; width and depth of
shaft at proximal end of intertrochlear foramen 26-1 X 6-2 ; length of foramen
5-2; maximum width of foramen 3-7 mm. Tibiotarsus. Greatest width across
anterior condyles at distal end 18-6; minimum width of posterior condyles at
distal end 15*3; greatest width of internal condyle 8-o; anterior length of
internal condyle 16-3; anterior/posterior width of external condyle 25-1;
width of shaft at proximal end of supratendinal bridge 13-6; width across
condyles at their proximal end on posterior surface 8-3 mm.
Cryptociconia, gen. nop.
Etymology. The generic name is formed from the Greek prefix crypto
(=hidden) and Ciconia (=a stork), in reference to the length of time during
which this form has been overlooked.
Type species. Cryptociconia indica, gen. et sp. nop.
Diagnosis. Distal end of tibiotarsus has large internal condyle with anterior
surface rounded but posterior tapering proximally to an abruptly terminating
flange. Internal ligamental prominence projecting only slightly. Anterior
portion of internal condyle narrow relative to its length.
Cryptociconia indica, sp. nop.
Etymology. The specific name refers to the country of origin.
Diagnosis. The only known species of its genus. Characters those of genus.
Material. An incomplete distal end of a tibiotarsus, B.M. (N.H.) no. 48444,
presented by Charles Falconer in 1868 (fig. 4).
Occurrence. Siwzlik Series, LowerPliocene;SiwalikHills,Uttar Pradesh, India.
Description. The holotype is a distal end of a left tibiotarsus, the shaft
broken off just proximal to the supratendinal bridge anteriorly; and to the
condylar groove posteriorly. The external condyle is extensively broken
anteriorly and posteriorly. Except for the narrowness of the anterior part of
the internal condyle the specimen shows a general similarity to the corres-
ponding parts of Recent Ciconia species in shape, but is considerably larger.
Viewed laterally, the internal condyle is rounded anteriorly, with a very
slight posterior slant, but posterior to this tapers towards the proximal
posterior end of the intercondylar groove where it fotms a prominent flange
terminating as an abrupt posterior projection. Anteriorly the smaller features
are damaged or eroded, but there is evidence of a laterally broad, medially
placed tubercle on the supratendinal bridge, the latter having a marked disto-
anterior slant. In length relative to width the anterior part of the internal
condyle is more similar to that of L. shvalicensis than those of the Recent
smaller storks which it otherwise resembles. In general size it would seem to
have been only slightly smaller than the large Recent Marabou, L. dubia.
Measurements. Greatest length of specimen 28-7; greatest distal width 20-7;
anterior/posterior thickness at internal condyle 30-3; posterior proximal
projecting flange 5-2; anterior length of internal condyle 17-8; thickness
(measured on outer side to condylar lip) 9-0; anterior/posterior thickness
at intercondylar groove 18-7; width of supratendinal bridge 7-6 mm.
SUMMARY
Various Pliocene and Miocene fossil specimens have been referred to the
extinct marabou stork, Leptoplilos falconer i. A re-examination reveals that
several species are involved. The distal end of the tibiotarsus, available for
all these species and often used as a holotype for extinct stork species, does
not show useful characters for separation in the present species although
such characters are present on other bones. It is suggested that the apparent
uniformity of a skeletal element such as this in birds in which other parts
differed might give rise to assumptions that Recent species existed at much
earlier periods when in fact morphologically distinct forms were involved.
Of eight Pliocene specimens originally referred to h. falconer i , five are here
considered referable to a very large species under this name; two to L.
shvalicensis (sp. nov.), a species of similar size to the Recent L. dubius; and one
to Cryptociconia indica {gen. et sp. nov.), an unrelated stork of similar size to the
last. The two Miocene specimens assigned by Rich to L. falconeri appear to
constitute another species, L. richae (sp. nop.).
References:
Brodkorb, P. 1963. Catalogue of fossil birds, part 1. Bull. Florida Sta. Mus., Biol. Set. 7: 180-
293.
— 1971- Origin and evolution of birds. In Farner, D. S. & King, J. R. (eds.). Avian Biol.
I: I 9 - 55-
Davies, W. 1880. On some fossil bird remains from the Siwalik Hills in the British Museum.
Geol. Mag. n.s. decade 2, 7: 18-27.
Lydekker, R. 1884. Siwalik birds. Mem. Geol. Surv. India 10: 3.
— 1 891. Catalogue of the fossil birds in the British Museum {Natural History). London: Brit.
Mus. (Nat. Hist.).
Milne-Edwards, A. 1 867-1 871. Kecherches anatomiques et pa/eontologiques pour servir a Vhistoire
des oiseaux fossiles de la France. Paris : Masson.
Rich, P. V. 1972. A fossil avifauna from the Upper Miocene Beglia Formation of Tunisia.
Notes Serv. Geol. Tunisia 35: 29-66.
The structure of feathers in Chlorophanes purpurascens
by L. Auber
Received 1 2th fanuary, 19J4
INTRODUCTION
The Trustees of the British Museum (Natural History) have been kind
enough to place at my disposal three contour feathers, each from a different
region of the plumage (chest, rump and crown), of the unique specimen of
Chlorophanes purpurascens Sclater & Salvin (1873, Nomen. Av. Neotrop.: 157.
Venezuela), redescribed and illustrated by Sclater (1886: 31 and pi. iv). This
specimen has also been considered by Hellmayr (1935 : 250). It is referred to
hereafter simply as "the holotype".
49
OBVERSE
CORTEX
INSERTION OF /&' (f
REVERSE
CORTEX
Figures from transverse sections through barbs from Chlorophanes purpurascens: — 1-4, in a
termino-basal succession from a chest feather (fig. 4 is a fragment of a section, showing
horizontal arrangement of medullary cells) ; 5 , through extreme base in vane of a rump
feather; 6-8, in a termino-basal succession through terminal vane half of a crown feather.
50
50,
Figures from transverse sections through broadest part of barb, all except 13 of a chest
feather: — 9, Chlorophanes spi^a <$; 10, Dacnis cay ana <$; 11, D. cay ana $; 12, Cyanerpes cyaneus
o* (from Auber 1957: fig. 8b); 13, C. cyaneus $ (of feather of interscapular region).
5i
Bond (in Phelps & Phelps 1948: 202-203) an< ^ Wetmore (in Storer 1957:
507) ascribe a hybrid origin to the holotype. They agree about one parent
species, Chlorophanes spi^a (the genus Chlorophanes being monotypic, if C.
purpurascens is not a distinct species), but disagree about the other one, Bond
believing it to be Dacnis cayana, Wetmore Cyanerpes cyaneus. Storer (op. cit.)
is inclined to agree with Wetmore, but does not entirely exclude the possi-
bility of it representing a valid species. All these putative parent species are
placed by Storer (1970: 391) in the subfamily Thraupinae of the family
Emberizidae.
On account of the great value of the material made available to me, for
many years I postponed sectioning the three feathers until I was sure that I
had a microtome knife of sufficient sharpness. Even so, the feathers from so
old a skin tended to splinter. Nevertheless they yielded the details described
below. Considering the probable hybrid origin of the holotype, detailed
discussion is necessary of the feather characters of the possible parent species.
INTACT SKIN AND FEATHERS
From my examination of the holotype as a whole, the purplish blue regions
of the plumage show a faint indication of an enamel effect, but no trace of the
glassy glitter peculiar to C spz\a. Under microscopical examination in
reflected light, of the intact blue barbs of the three feathers available, the
reverse surfaces all show a black longitudinal zone. In transmitted light the
same barbs show, outside a black opaque zone, one that transmits yellowish
light (the medulla, see fig 1), and adjacent to this a transparent colourless
zone belonging to the cortex. Evidently the blue colour of these feathers is
due to the optical phenomenon that causes most instances of non-iridescent
blue, green and purple in feathers. After mistakes by previous physicists, this
optical effect has been recognised by Dyck (1971) as a special instance of
interference of light, and is not a Tyndall effect.
TRANSVERSE SECTIONS THROUGH BARBS (figS. 1-8)
When stained according to the routine method of Auber (1955), the
medullae did not attain the typical acid stain with picro-Indigocarmine, but a
pale orange stain due to an apparently not very strong affinity with both
Basic Fuchsin and picric acid. Owing to the restricted amount of material,
it is not possible to explain this unusual staining reaction.
Cortex: The obverse cortex (figs. 1-3) is rather massive and laterally com-
pressed into an approximately prismatic shape with a fairly blunt summit. In
routine-stained sections it appears practically homogeneous, except that
the marginal portions (corresponding to surface cells) have less affinity with
Basic Fuchsin than have the portions inside the prism. The reverse cortex
has an intense melanisation which extends laterally up to the likewise
melanised barbules. Towards the concealed vane portions (fig. 5) of the
feathers of the chest and rump, the obverse cortex is further compressed into
a narrow ridge, so that the outline is pear-shaped. Melanisation in the reverse
and lateral cortex is reduced.
Medulla: On the chest and rump, near the attenuated barb tip (fig. 1), the
medulla has a vertically oval outline. Around the widest portion of the barb
(figs. 2-3) the outline is almost circular, the cells having almost a radial
arrangement. Occasionally, as in the fragment (fig. 4), the cells form a hori-
zontal series. However, this arrangement has been found only near the vane
base, although apparently still in the exposed feather portion. Throughout
the medulla (figs. 2-4) the cell-boundaries are fairly straight. Each cell
contains a cavity of a rather irregular shape, often with rounded ("ameboid")
diverticles. Their rounded outlines indicate that these shapes ar e not artefacts.
More jagged outlines would result, if the cell walls splintered during sec-
tioning. In the chest and rump feathers the medulla is altogether amelanotic.
In the blue terminal half of the crown feathers (figs. 6-8) nearly every cell of
the laterally compressed medulla contains small black granules of rodlet-
shape, irregularly distributed in varying amounts.
DISCUSSION
Hellmayr (1935 : 250) considered the holotype to be a male. This conclusion
is apparently based on the chiefly blue shade of the plumage. In most coerebo-
thraupines the males are mainly blue, the females mainly green (due to
yellow filter pigment being superimposed on the structural blue). Bond's
opinion (in Phelps & Phelps 1948) that the holotype is a hybrid between
Chlorophanes spi^a and Dacnis cayana is based on: — (1) the overall similarity
in shape with C. spi^a; (2) the similarity in main colour-pattern with D.
cayana (^); and (3) that it is partly sympatric with these two species. Storer
(1957) believes that Wetmore, whom he quotes, is more correct in assuming
that the parent species are Chlorophanes sp/\a and Cyanerpes cyaneus (not D.
cayana). The following reasons are given: — (1) there is a greater similarity
in the colour-pattern between the holotype and Cyanerpes cyaneus (<^), and
likewise in the dark purplish main colour (light blue in D. cayana $); (2) the
bill of the holotype, quite similar to that of Chlorophanes spi^a, is more
similar to that of Cyanerpes than to that of Dacnis. At all events, the sympatric
argument made by Bond applies also to Cyanerpes cyaneus.
Feather structure: Individual feather characters in the holotype (figs. 1-8)
are dealt with under (a) - (f) below. Items (a) and (b), and to some extent (c),
indicate affinity with Chlorophanes spi-^a, but none of them indicate any special
affinity with either Dacnis or Cyanerpes. These items may now be detailed:—
(a) Prismatic obverse cortex: Approaches the conditions in Chlorophanes
spi^a <$ (cf. fig. 9), but is much lower and lacks the extreme sharpness of its
summit (height and sharpness of summit, together with absence of barbules
from exposed feather portions, causing glitter in C. sp/\a $). Neither does
the prism in the holotype contain the complex light-scattering apparatus
described by Auber & Appleyard (1955 : 255-256) which in the C. spi^a <$
imparts a green shade to the structural blue. This atypical cause of the green
is an alternative to the usual cause, viz. superposition of yellow filter pigment
on blue-producing structures : see below under females. In this context male
skins in the Naturhistorisches Museum, Vienna, mislabelled 'Chlorophanes
purpurascens\ without further data (cf. Auber & Appleyard 1955) are in
reality C. spi\a with a strong bluish tinge to the green. According to a recent
inquiry these specimens were collected near Quito, and so can be determined
as C. s. caerulescens (cf. Sclater 1886: 30). The bluish tinge of these specimens,
as compared with the intense green of eastern C. spi^a <$$, can be attributed
to a less efficient light-scattering apparatus in the prism. In Dacnis cayana £
(fig. 10) and the much more highly specialised Cyanerpes cyaneus £ (fig. 12) the
obverse cortex forms a very thin lamina.
(b) Melanisation of reverse cortex (configuration [Ba] in Auber 1957: 462):
Agrees with the condition in both sexes of Chlorophanes spi^a (fig. 9) except
that: — (1) in the holotype the melanisation extends into well developed
barbules (such being absent from the exposed green vane portions in C.
spi^a ^, whereas corresponding portions in C. spi^a $ bear amelanotic
barbules); (2) melanisation at the barb bases is rarefied (fig. 5), whereas in
53
C. spi^a fairly intense melanisation of the reverse cortex coexists with
melanisation in the extreme obverse (Auber 1957: fig. 15 c!). In both sexes of
Dacnis (figs. 10, 11) and in Cyanerpes ft (fig. 12) melanisation is contained in a
cortical partition (configuration [Be] in Auber 1957: 462).
(c) Irregular melanisation of medullary cells in crown feathers (figs. 6-8): The
first impression given by this character is of an extreme instance of an other-
wise sporadic occurrence in typically amelanotic medullary cells of various
bird forms. The crown feather of the holotype is not melanised to such an
extent that it appears entirely black in situ (as do corresponding feathers in
Chlorophanes spi^a ft). However, it approaches the condition in the head
region of this speciqs and sex, contrary to that in the chest and ramp feathers
of the holotype and in any blue feather from Dacnis and Cyanerpes ftft, none
of which show any black in situ.
(d) Pear-shape of sections (fig. 4) through barb bases is a generalised feature,
indicated in at least the vane base of most feather types (Auber 1957: 460-
461).
(e) Medulla of oval or circular outline (figs. 1-3, 5): Again a generalised
character. It is absent from exposed barb portions in structurally coloured
feathers from both sexes of Chlorophanes spi^a (fig. 9), of Dacnis (figs. 10, 11)
and from Cyanerpes ft (fig. 12): in all these instances the medullary cells form
horizontal tiers. Thus in the holotype the apparently occasional horizontal
arrangement (fig. 4) of medullary cells is not significant in the present
context.
(f) Irregular {partly "ameboid") shape of medullary cell cavities: As far as could
be seen, peculiar to the holotype. It contrasts with an entire absence of
medullary cavities from exposed barb portions in structurally coloured
feathers of Chlorophanes spi^a (fig. 9), as well as with the condition in Dacnis
and Cyanerpes (figs. 10-13), in which these barb portions contain medullary
cells with cavities of regular "polyhedral" shape (cf. Auber 1973). Absence of
cavities from medullary cells in Chlorophanes spi^a appears to be unique (cf.
Auber & Appleyard 1955: 253, 255). Polyhedral cavities, on the other hand,
are frequent in Passeres, but they also occur in some non-passerine families.
Females: In those of all three possible parent species the plumages are
chiefly green, viz.: — (1) in a typical fashion, yellow filter pigment in the
transparent obverse cortex is superposed on the blue-producing structure
(cf. Kniesche 19 14); (2) often also, green barbs differ from corresponding
blue barbs by a more massive obverse cortex (Frank 1939: 501-502), to
produce a more efficient filter. Character (2) applies to the bright green in
Dacnis (cf. fig. 10 with fig. 11), in which form the blue and green barbs of
males and females, respectively, otherwise show the same level of speciali-
sation. This character also applies to the dull green in Cyanerpes .$. In this
form, however, the ft (fig. 1 2) is much more highly specialised than are both
sexes of Dacnis. The Cyanerpes $ (fig. 13), on the other hand, shows structu-
rally some generalised details (sections through barbs pear-shaped and
melanisation confined to barbules, cf. configuration (C) in Auber 1957: 462).
In females of the extremely highly specialised Chlorophanes spi^a only charac-
ter (1) obtains: the obverse cortex forms a massive prism in both sexes.
SUMMARY
Recent authors interpret the unique specimen originally described as
Chlorophanes purpurascens as a hybrid between two coerebo-thraupine species,
either Chlorophanes spi^a and Dacnis cayana or C spi^a and Cyanerpes cyaneus.
Examination of structurally coloured feathers from C purpurascens shows
54
that they agree in certain respects with those of C. spi^a but do not show the
extreme degree of specialisation found in that species. Along their barbs they
show considerable fluctuation, in details of structure and melanisation. On
the other hand, no details could be found pointing towards the structures in
the feathers of the other species suggested as a parent, either Dacnis cayana or
Cjanerpes cyaneus.
AC KNOWLEDGEMENTS
My thanks are due to the Royal Scottish Museum for facilities for research;
to the Trustees of the British Museum (Natural History) and Mr. I. C. J.
Galbraith for the unique material; to Mr. Galbraith also, and Dr. A. S.
Clarke, Mr. I. H. J. Lyster, Mr. J. M. Sanderson and Mr. A. R. Waterston
for constructive criticism; to Dr. G. von Rokitansky and Prof. R. W.
Storer for valuable information; to Mr. C. W. Benson for assistance in
presentation of this paper; to the Royal Society for a research grant; and to
the Editor, Journal of Zoology, for permission to reproduce the present fig.
12.
References :
Auber, L. 1955. Cortex and medulla of bird-feathers. Nature 176: 1218-1219.
— 1957. The structures producing "non-iridescent" blue colour in bird feathers. Proc. Zool.
Soc. Fond. 129: 455-486.
— 1973. Formation of "polyhedral" cell cavities in cloudy media of bird feathers. Proc.
Roy. Soc. Edin. (B) 74(2): 27-41.
Auber, L. & Appleyard, H. M. 1955. The structure of the feathers in Chlorophanes and
Iridophanes (Coerebidae). Ibis 97: 252-258.
Dyck, J. 1 97 1. Structure and colour production of the blue barbs of Agapornis roseicollis and
Cotinga maynana. Z. Zellforsch. 115: 17-29.
Frank, F. 1939. Die Farbung der Vogelfeder durch Pigment und Struktur./ow«. Ornith. 87:
426-523.
Hellmayr, C. E. 1935. Catalogue of birds of the Americas and the adjacent islands. Field
Mus. Nat. Hist., Zool. 13(8) (publ. 347): 541 pp.
Kniesche, G. 1914. Uber die Farben der Vogelfedern. I. Die Griinfarbung auf Grundlage
der Blaustruktur. Zool. Jahrb. (Anat.) 38: 327-356.
Phelps, W. H. & Phelps, W. H., Jr. 1948. Contribucion ornitologica de la coleccion
Phelps, 5 : Notes sobre aves Venezolanas. Bol. Soc. Vene^ol. Cienc. Nat. 10(72): 189-210.
Sclater, P. L. 1886. Catalogue of the birds in the British Museum 1 1. London: Brit. Mus. (Nat.
Hist.).
Storer, R. W. 1957. The hybrid origin of Chlorophanes purpurascens . Auk 74: 507.
— 1970. Subfamily Thraupinae, pp. 246-408 in Check-list of birds of the world 13, R. A.
Paynter, Jr., ed. Cambridge, Mass. : Mus. Comp. Zool.
The type of the
Madagascar Cuckoo Cuculus poliocephalus rochii Hartlaub
by C. W. Benson
Received 2jth February, 1974
Benson el al. (1970: 17) considered that a female of this cuckoo in the
University Museum of Zoology, Cambridge, collected at Chasmanna,
Madagascar, on 2nd October 1862, could not be the type, even although it
had been indicated as such by Prof. A. Newton. This was because Hartlaub's
description (1862: 224) was ostensibly published on 11 November 1862, only
40 days after the date of collection. However, Duncan (1937: 72) gives the
true date of publication as April 1863. Thus there was a lapse, not of a mere
40 days, but of some six months, so that even in those more leisurely times
completion of the process from collection to publication of a description
would have been possible.
55
What must be the specimen in question is referred to by E. Newton (1863:
453). He states that it would soon have bred, while according to the label it
had "eggs large". Roch & Newton (1863: 166) refer to a specimen which
might be supposed to have been collected in September or October 1861
{ibid., 1 862 : 265-266). No such specimen has been found in either Cambridge
or Bremen, where there are many of Hartlaub's types (Benson eta/. 1970 : 17).
Reference to the Ibis for 1863 shows that the second part of Roch & Newton's
paper was published in April of that year. The note on Cuculus rochii (p. 166)
contains a reference to Hartlaub (1862), presumably a last minute insertion,
with the "One obtained" referring to the specimen available to Hartlaub.
The matter has been discussed with R. Wagstaffe, who agrees that the
Chasmanna specimen may be properly regarded as the holotype, contra also
Benson (1971: 4), who does list several other Hartlaub Malagasy types in
Cambridge. Incidentally, Benson et al. (1970: 17) suggest that the type of
rochii was collected somewhere between Tamatave and Tananarive. Chas-
manna has not been traced from any map or gazetter, either modern or from
the last century. However, from the account of his travels during his second
visit to Madagascar (E. Newton 1863: 333-335), Chasmanna cannot have
been far upstream from the mouth of the Hivondrona (now Ivondro) River,
just to the south of Tamatave. For practical purposes the type locality may be
taken to be Tamatave.
I gladly absolve my co-authors in Benson et al. (1970) from responsibility
for any previous error, more particularly the overlooking of the paper by
Duncan (1937).
References :
Benson, C. W. 1970-71. The Cambridge collection from the Malagasy Region. Bull. Brit.
Orn. CI. 90: 168-172; 91: 1-7.
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. Stuart. 1970. Notes on the
birds of Zambia. Arnoldia {Rhodesia) 4(40): 59 pp.
Duncan, F. M. 1937. On the dates of publication cf the Society's "Proceedings", 1859—
1926. Proc. Zool. Soc. Lond. 107A: 71-84.
Hartlaub, G. 1862. On a new bird from the island of Madagascar. Proc. Zool. Soc. Lond. 31 :
224.
Newton, E. 1863. Notes of a second visit to Madagascar. Ibis 5 : 333-350, 452-461.
Roch, S. & Newton, E. 1862-63. Notes on birds observed in Madagascar. Ibis 4: 265-275 ;
5: 165-177.
The Pintail Anas acuta in Rhodesia
by Michael P. Stuart Irwin
Received 1st March, 19 J 4
The holarctic Pintail, Anas acuta Linnaeus, winters extensively in northern
tropical Africa (Moreau 1972 : 220), although it has been collected once as far
south as Zambia at Ndola at 13 S., in February (Benson et al. 1971: 55).
Within the South African sub-region the only evidence of its occurrence is
provided by Took (1959: 84) who observed a pair near Salisbury in Novem-
ber 1958.
On 3rd February 1974 Mr. Eugene Halsted, while on a shooting party,
obtained two female Pintails from a flock of five on a farm dam at Portwe
Estate, i9°4o'S., 28°42'E., near Turk Mine. Both carried extensive body fat
and weighed 822-5 an d 852-7 grammes respectively. During the following
week a flock of six were reported from the same locality, by Mr. Huntsman
Williams who had seen the original group of five and had become familiar
with the species. Garganey Anas querquedula Linnaeus were also reported as
56
having been shot at the same time as the Pintail, but no specimens were
preserved.
It is therefore possible that an influx of these palaearctic duck may have
occurred. Exceptionally heavy rains have characterised the 1973/74 rainy
season in Rhodesia and over most of south-central Africa. This factor,
accompanied by the progressive series of droughts in the northern semi-
arid savanna belt of Africa (Winstanley 1973 : 190-194), may well force many
birds south of their present normal wintering area. That Pintail, at least
during the cooler periods of the Pleistocene, must have occurred very much
further south than they do to-day, is evidenced by the existence of isolated
resident, subspecifically distinct, sub-antarctic populations found on Ker-
guelen and the Crozet Islands (Delacour 1956: 132-135).
My sincere thanks are due to Mr. Eugene Halsted for reporting the pre-
sence of these ducks, and to Mr. Robert Halsted for bringing this material
into the National Museum, Bulawayo, for identification.
References :
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. S. 1971. The birds of Zambia.
London: Collins.
Delacour, J. 1956. The waterfowl of the World, vol. 2. London: Country Life.
Moreau, R. E. 1972. The Palaearctic- African bird migration systems. London & New York:
Academic Press.
Took, J. M. E. 1959. Pintail Anas acuta in Southern Rhodesia. Ostrich 30(2): 84.
Winstanley, D. 1973. Rainfall patterns and general atmospheric circulation. Nature 245
(no. 5422): 190-194.
[R. J. Dowsett {in litt.) reports the following records of palaearctic Anatidae from Zambia
in the 1973-74 rains: — Pintail: Kitwe, one 26 Dec. to 3 Jan. (B. Coates, Dr. E. H. Penry);
Mufulira, one 8 to 13 Jan. (T. Sims, G. P. Robinson). European Shoveler Anas clypeata:
Kitwe, four 25 Dec, three 27 Dec. and 1 Jan., one 17 Jan. (Coates, Penry). Garganey Anas
querquedula: Recorded from Mufulira, Kitwe, Blue Lagoon and Lochinvar, first at Mufulira
30 Nov. Maximum numbers (the largest yet recorded from Zambia) were at Lochinvar in
Mar., when 100+ were counted (T. O. Osborne, R. Stjernstedt). — Ed.]
Larus tninutus in Angola
by J. C. Sinclair
Received 10th December, iyj)
A very small gull was sighted flying overhead in Lobito harbour, Angola,
ca. i2°S., on 6 May 1973. The bird was seen again a few minutes later, when
sufficiently adequate views were obtained to identify it as a Little Gull Larus
minutus Pallas, and the following description was noted: — Upperparts: fore-
head and front of crown white; rest of crown, nape, mantle and wings grey;
leading edges of primaries black, likewise edges of secondary coverts which
formed a conspicuous W on upper- wings ; square-ended tail white narrowly
tipped black. Underparts: — mainly white with pale underwings.
The bird was in company with Black Terns Chlidonias nigra, and was
noticeably larger. The flight was bouyant and tern-like, on wings which had
a rounded appearance at the tip. The bill was short, stubby and dark in
colour. No black spot behind the eye was apparent.
This record appears to be the first for the Little Gull south of the Canary
Islands (Watson 1965, Seabirds of the tropical Atlantic Ocean), and the species
is not mentioned by White (1965, A revised check-list of African non-passerine
birds).
[The above note has been read by R. Wagstaffe. With long experience of palearctic gulls,
he considers the record authentic. — Ed.]
57
Fish offal scavengers off Luanda
by J. C. Sinclair
Received ioth December, iyjj
Brooke (1971, Bull. Brit. Orn. CI. 91: 46) records Little Egrets Egretta
gar^etta and Grey Herons Ardea cinerea scavenging inside the harbour at
Luanda, Angola. On 10 and 1 1 May 1973 dawn sea- watches were carried out
at the northern tip of the "island" just off Luanda. The "island" runs parallel
to the coast and forms the outer bar of the bay, the northern tip being at the
bay entrance. During the course of the morning many small fishing craft
entered the bay, gutting fish and throwing the offal overboard. Groups of
ca. 500 Little Egrets and ca. 140 Grey Herons were assembled on the sandy
point not far from the Casuarina trees where they breed and on sighting a
boat offshore would fly out to meet it and follow it into the bay entrance
picking up, in the manner described by Brooke {pp. cit.), pieces of offal
discarded by the boats. The Little Egrets were noticeably more adept at
foraging than the ponderous Grey Herons. They would even take fish out of
unattended baskets when the boats had been berthed. Some boats were joined
by the herons and egrets at a distance of up to a mile offshore. A Grey Heron
was seen to plunge and settle on the water to peck at a large piece of offal and
had no great difficulty flying up and pursuing the boat again. Other species
seen following the boats were one immature black-backed gull either Larus
fuscus or dominicanus, Arctic Skuas S tenor arius parasiticus (one skua made an
attempt to harry a Little Egret but with little success), Royal Terns Thalasseus
maximus, Black Terns Chlidonias nigra and a Palm-nut vulture Gypohierax
angolensis. The last named, although it did not directly follow the boats, did
pick a fish from the water surface in its talons.
I am obliged to R. K. Brooke for commenting on a draft of this note.
Vocal mimesis in nestling Greater Honey-guides
by C. H. Fry
Received iyth December, 1973
In February and March 1973 I studied the social organisation of Red-throated
Bee-eaters Merops bulocki in breeding colonies at Zaria, n°o8'N, 7°47'E,
northern Nigeria. Using an industrial endoscope (fibroxscope) with portable
6v light source, I was able to make regular observations not only of young bee-
eaters in situ in their nests at the end of 1 m long tunnels, but also of young
Greater Honey-guides Indicator indicator that the bee-eaters fostered.
African honey-guides parasitise a wide variety of hole-nesting birds, with
some preference by Indicator minor and /. variegatus for other piriform species,
particularly barbets (Friedmann 1955, Proc. U.S. Nat. Mus. 108: 309). Al-
though /. indicator often parasitises barbets and woodpeckers, a much greater
number of records involve coraciiform hosts, particularly six or more Merops
species which constitute about one third of the known cases (Friedmann 1970,
Los Angeles Co. Mus. Contrib. Sci. 205 : 1). Most tropical bee-eaters lay 2-4
eggs; many of these are destroyed by the female honey-guide, and young bee-
eaters hatching from unbroken or subsequently laid eggs are, in most cases,
destroyed by the newly-hatched honey-guide, which usually comes to occupy
the nest alone.
Nestling M. bulocki have a loud and insistent food-soliciting call "sif-sif-sif-
sif . . .", about five syllables per second. At least until their eyes begin to open
at about eight days, it is easily elicited by a variety of stimuli, with the intensity
of which its loudness and frequency are positively correlated. Using the endo-
scope for visual corroboration, I found it easy to distinguish by ear a nestling
calling alone from more than one nestling calling simultaneously, but very
difficult to distinguish two from three or more.
The begging call of the nestling L indicator is a loud and insistent, irregularly-
timed series of "sifs", 5-10 syllables per second. It is delivered with the head
sunk into the shoulders and the beak held vertically upwards. Skead (195 1,
Auk 68 : 52) described the call as "husky and perpetual". He likened it to the
guiding call of the adult /. indicator and noted that it can be heard, greatly
accelerated and intensified, 75 yards from the nest whenever a foster parent
approaches with food. To my ear the voice was not so much husky as sibilant,
and extremely similar to the voice of nestling M. bulocki. However, the food-
soliciting call of the nestling /. indicator resembles not one but two (or more) M.
bulocki nestlings calling simultaneously.
Vocal convergence between parasite and host nestlings is not unexpected,
in view of the diversity of mimetic resemblances that has evolved between
brood parasites and their hosts at all growth stages (Lack 1968, Ecological
adaptations for breeding in birds). Indeed such convergence has been suggested
for some cuckoos (Courtney 1967, Emu 67: 1 54-157). But the call of a nestling
/. indicator is likely to evoke the feeding response in an adult bee-eater more
powerfully than the call of a single bee-eater nestling. If so, it would confer
the selective advantage that the parasite would compete effectively for the
attention of provisioning adults with neighbouring nests in the colony, however
many nestlings they contain. In this and other colonial species of Merops,
young in the nest are fed not only by their parents but by adult non-breeding
helpers whose bonds with the parents are not strong enough to ensure their
complete fidelity to their adopted nest (Fry 1972, Ibis 1 14: 1). Even were there
no helpers, the honey-guide call is probably a super-normal stimulus that
elicits the feeding response of any passing adult bee-eater, just as the begging
call and wide, brightly coloured gape of the young Cuckoo Cuculus canorus does
with adult passerines. Observations at one parasitised nest on 16 March
suggested that during a three hour period at least four adult bee-eaters fed
the 1 5 -day old honey-guide chick.
At least up to two weeks of age, /. indicator nestlings were found to be
extremely voracious; in the nest they tried to engulf the 10 mm diameter tip
of the endosope, and when handled one bird attempted to swallow a finger.
Why provisioning bee-eaters are not harmed by the nestling's needle sharp
mandibular hooks is not at all apparent.
I am most grateful to Dr. H. Friedmann and Dr. J. B. Nelson for their
comments upon an earlier draft of this paper.
Buteo tachardus Andrew Smith 1830
by R. K. Brooke
Received 21st November, 19J3
Roberts (1940), Courtenay-Latimer (1941) and Clancey (195 1) showed that
South Africa had a small resident Buteo as well as the well known large B.
rufofuscus (Forster) 1798. These authors associated it with B. oreophilus
Hartert & Neumann 1914 with type locality Koricha in Ethiopia. Rudebeck
(1957) demonstrated that the small South African Buteo was not the same as
the northeast African B. oreophilus and proposed for it the name Buteo buteo
59
tri^onatus with type locality Knysna in the southern Cape Province of South
Africa. He described it as smaller and duller than B. b. vulpinus (Gloger) 1833
and with clear white between the brown streaky markings of the breast and
lower abdomen. However, although he was unaware of it, he was not the
first author to describe and name this bird. This was done by Smith (1830:
381) who in discussing the Buteo species of what is now the Cape Province
listed B. lagopus by which he meant what we know as Hieraaetus pennatus
(Gmelin) 1788 (Roberts 1936 thought so and I concur), B. tachardus by which
he meant what we have known as B. b. triyonatus Rudebeck 1957 and B.
desertorum (Daudin) 1800 which is an indeterminable name normally applied
in the past, as here, to what we know as B. b. vulpinus (Gloger) 1833.
Smith (1830), as many others have done, found Buteo specimens confusing.
Nonetheless, he drew attention to and named a small form (18 J ins. long,
i.e. just over 460 mm) with a clear white upper abdomen between the brown
streaked and blotched breast and lower abdomen. The significant part of his
description reads: —
"under parts white, with the throat streaked by narrow longitudinal
brown lines, and the breast and posterior (my italics) part of the belly
more or less spotted with oblong or roundish brown blotches".
The description is repeated with slightly different wording and punctuation
in Smith (1834). He believed that he was dealing with Le Vaillant's (1799) Le
Tachard and Falco tachardus Daudin 1800, both of which are synonyms of
Perms apivorus (L.) 1758 (Hartert 1914). What Smith (1830) thought he was
doing is of little importance in comparison with what he did do, i.e. describe
a taxon in Buteo which subsequent study has shown to be valid : cf. Vaurie
(195 1) on the validation of Hirundo lagopoda Pallas 181 1 where the latter also
confused the issue with irrelevant citations. Two of Smith's specimens of B.
tachardus were examined in the British Museum (Natural History) by Rude-
beck (1958) who placed them with his tri^onatus so there can be no doubt that
the form was known to Smith.
I conclude that Buteo tachardus Smith 1830 is an unequivocal description of
what Rudebeck (1957, 1958) clarified under the name B. b. tri^onatus and that
since Smith's name has undisputable priority and does not fall within the
conserving terms of Articles 23 and 79 of the International Code of Zoo-
logical Nomenclature as set out in Bull. Zool. Nomencl. 29: 185-186, it must
be used by any zoologist who believes that the small South African Buteo is
an objective taxon.
Rudebeck (1957) gave the type locality of his tri^onatus as Knysna, Cape
Province, South Africa. Since by 1830 Smith had not worked outside what
is now the Cape Province (Roberts 1936), Knysna can be regarded as the
restricted type locality of B. tachardus. Smith's surviving specimens of this
form are labelled only as coming from South Africa (Rudebeck 1958) but
there is no reason why they should not have been obtained at or near Knysna.
If a lectotype is required, one of these specimens should be chosen.
Following Siegfried (1971) I regard B. tachardus Smith 1830 and B. oreo-
philus Hartert & Neumann 19 14 as conspecific and not part of B. buteo (L.)
1758. The new nomenclatural combinations required are, therefore, Buteo
tachardus tachardus and Buteo tachardus oreophilus with Buteo buteo tri^onatus as a
subjective synonym of B. t. tachardus. For check list purposes the name
should appear as : —
Buteo tachardus tachardus Smith
Buteo tachardus Smith, 1830, S. Afr. Quart. Journ. 4: 381. South Africa,
60
restricted to Knysna, Cape Province, South Africa by Rudebeck (1957,
S. Afr. Anim. Life 4: 418).
I have examined the type of B. b. tri^onatus Rudebeck and all the specimens
in South African museums that he placed with it (Table 1) and concur in his
comments (Rudebeck 1957, 1958) as far as they go. I have not found any
other specimens which should be referred to this form. A study of the moult
of the primaries on the lines of Brooke et al. (1972) has elucidated certain
TABLE 1
Measurements in mm of Buteo t. tachardus specimens.
use urn and no.
Type
Age
Sex
Wing
Cnlmen
Hind clarv
Museum and no.
ad.
c?
340
21
19
T.M.P. 6278
ad.
cJ
340
21
19
24273
ad.
?
345
23
22
E.L.M. 578
ad.
?
350
23
23
N.M.PMB —
ad.
oj?
350
21
20
T.M.P. 61
ad.
o«J?
335
21
19
A.M.G. 968
imm.
oJ?
330
21
21
T.M.P. 6281
imm.
?
350
21
22
6280
imm.
0??
365
23
21
62
imm.
o$?
350
22
23
6279
juv.
?<??
335
20
19
545°
juv.
0??
340
22
—
„ 5006
aspects of plumage succession. On the apparently well founded assumption
that juveniles (first year birds) show no moult of the primaries, immatures
(second year birds) show a simple descending mode of moult and adults a
jumbled mode of moult (see also Stresemann & Stresemann 1966 on B. b.
buteo) it can be stated that the juvenile and immature plumages of B. t.
tachardus are characterized by leg feathers which are white with a few brown
spots or blotches (this is not individual variation as believed by Smith 1830)
whereas in adults and in all age classes of B. b. vulpinus the legs are predomin-
antly dark brown of some shade. In addition, the darkest specimens of
tachardus are all adults but the sample I have seen (Table 1) is too small to say
if this is always true in a member of a genus as notoriously variable as Buteo
is.
Although Siegfried (1971) accepted the sight record of B. t. tachardus in the
northern Potgietersrus District of the Transvaal (not far from The Downs) by
van der Merwe & Pienaar (1959), I do not. The latter authors give no details
in support of their record of an apparent rarity; they do not record the much
more probable B. b. vulpinus despite visiting the area in summer (judging by
the number of summer visitors they list : they do not give the dates of their
visits) ; they record for the area a number of species which are not known
from anywhere near Potgietersrus, e.g. Turdoides leucopygius, Lanioturdus
torquatus, Onychognathus nabouroup, again without comment.
I am obliged to Mr. C. Jacot-Guillarmod of the Albany Museum in
Grahamstown, Dr. A. C. Kemp of the Transvaal Museum in Pretoria, Dr.
J. A. Pringle of the Natal Museum in Pietermaritzburg and Mr. C. Quickel-
berge of the East London Museum for facilities for study and to Mr. P. A.
Clancey for commenting on a draft of this paper.
References :
Brooke, R. K., Grobler, J. H., Irwin, M. P. S. & Steyn, P. 1972. A study of the migratory
eagles Aquila nipalensis and A. pomarina (Aves: Accipitridae) in southern Africa, with
comparative notes on other large raptors. Occ. Pap. Nat. Mus. Rhod. B5 : 61-114.
Clancey, P. A. 195 1. Notes on the birds of the South African sub-continent. Ann. Natal
Mus. 12: 137-152.
61
Courtenay-Latimer, M. 1941. Breeding of the Mountain Buzzard. Ostrich 12: 20-23.
Hartert, E. 1914. Die Vogel der pal darktischen Fauna II. Berlin.
Le Vaillant, F. 1799. Histoire naturelle des oiseaux d'Afrique I. Paris.
Roberts, A. 1936. Some unpublished field notes made by Dr. (Sir) Andrew Smith. Ann.
Transv. Mus. 18: 271-323.
— 1940. The birds of South Africa. London : H. F. & G. Witherby.
Rudebeck, G. 1957. Buteo buteo tri^pnatus, a new buzzard from the Union of South Africa.
S. Afr. Anim. Life 4: 415-437.
— 1958. Some additional notes on the buzzard, Buteo buteo tri^pnatus. Bull. Brit. Orn. CI. 78 :
54-56.
Siegfried, W. R. 1971. Affinities of the small African and palaearctic buteos. Ostrich Suppl.
8: 41-46.
Smith, A. 1830. A description of the birds inhabiting the south of Africa. S. Afr. Quart.
fourn. 4: 380-391, per Willughby Soc. reprint.
Smith, A. 1834. African zoology. S. Afr. Quart. Journ. 273-288, per Willughby Soc. reprint.
Stresemann, E. & Stresemann, V. 1966. Die Mauser der Vogel. fourn. Ornith. Sonderheft.
van der Merwe, N. J. & Pienaar, D. 1959. A preliminary list of the birds of the Percy Fyfe
Nature Reserve. Ostrich 30: 130-133.
Vaurie, C. 1951. Notes on some Asiatic swallows. Amer. Mus. Novit. 1529.
The African southern limits of the Steppe Eagle in winter
by R. K. Brooke
Received 21 st January, 1974
Brooke, Grobler, Irwin & Steyn (1972, Occ. Pap. Nat. Mus. Rhod. B5 : 61-
114) set out the known records of the Steppe Eagle Aquila nipalensis orientalis
in southern Africa in detail and predicted that, at least at times, it should
extend south into the Karoo of the Cape Province of South Africa since its
predominant prey, alates of the Harvester Termite Hodotermes mossambicus,
are abundant there. This proves to be the case. The East London Museum
has a specimen received from J. Pete, a farmer in the Tarkastad District of
the Cape Province, who used to trap birds for his aviaries and pass on to the
museum those that died. Hence the date of death of this specimen, 1 5 June
1942, has nothing to do with its time of occurrence near Tarkastad other than
to provide a terminus post quern. The specimen is a very dark fourth year male
on the characters set out in Brooke et al. {pp. cit.) with wing 585, culmen from
cere 41, hind claw 32 mm. It may be assumed that it was trapped on or close
to fete's farm, i.e. ca. 32°io / S, 26°oo'E, as were his other birds (Dr. M.
Courtenay Latimer pers. comm.).
Clancey (195 1, Ann. Natal Mus. 12: 137-15 1) reported a specimen of
A. n. orientalis from Himeville, Natal at 29^5 'S, 29°3i'E. Upon examination
of the specimen in the Natal Museum, Pietermaritzburg, I find it to be a first
year male with wing 575, culmen from cere 36, hind claw 32 mm.
I am obliged to Dr. M. Courtenay-Latimer, Director of the East London
Museum, and to Dr. J. A. Pringle, Director of the Natal Museum, for facili-
ties for study.
A further record of the White-eyed Gull
Icarus leucophthalmus from Beira, Mozambique
by B. G. Donnelly
Received 19th fanuary, 1974
On 12 March 1972, a small gull fitting the description of a second winter or
non-nuptial plumaged White-eyed Gull Larus leucophthalmus (see Dwight
1925, Bull. Amer. Mus. Nat. Hist. 52: 152) was observed at the fishing
62
harbour at Beira. The bird was feeding with numerous Grey-headed Gulls
L. cirrocephalus, and a single Lesser Black-backed Gull Larus fuscus. In flight
the bird was more agile than L. cirrocephalus and slightly smaller in size as
mentioned by Liversidge (1964, Ostrich 35 : 111-112). The overall plumage
colour was bumsh-white while the tail had a sub-terminal band of dark spots.
The head was greyish with an imperfectly formed black hood. The bill, legs
and feet were yellowish-brown. The dark primaries and secondaries formed
a conspicuous terminal band on the wings.
The only other sightings of this species from southern Africa were made
at Beira (Benson 1948, Ostrich 19: 1 50-1 51) and Port Elizabeth (Liversidge
op. cit.), both during the month of January. These and the present record
suggest that individuals of this species may only move this far south of their
usual range around the Red Sea and Gulf of Aden (Archer & Godman 1937,
Birds of British Somali land and the Gulf of Aden 2) during December to March.
Incidentally, Liversidge was dubious about Benson's record, but Benson
(pers. comm., 25 July 1973) states that the bird which he saw was most
carefully identified with the aid of Alexander (1928, Birds of the Ocean.
London: Putnam).
The Lesser Black-backed Gull Larus fuscus
in southern and central Africa
by B. G. Donnelly
Received 19th January, 1974
Only recently has the occurrence of Larus fuscus in southern Africa been
generally accepted (McLachlan & Liversidge 1970), while data on this
species from central Africa are scarce. This communication brings together
the published data and several new observations, and discusses birds pre-
viously identified as L. dominicanus.
Larus fuscus breeds in Iceland, the Faeroes, along the coasts of the British
Isles, Scandinavia, Finland and the Baltic and winters on the west coast of
Africa, the Mediterranean, Red Sea and the Gulf of Aden with some birds,
L. f fuscus, moving inland to the Great Lakes of East Africa (Archer &
Godman 1937; Vaurie 1965). Moreau (1972) points out that L. fuscus is often
abundant at times in East Africa and that they range as far south as Botswana
around 18 S.
The closely related Southern Black-backed Gull L. dominicanus occurs
around the coast of southern Africa from Angola to Lourenco Marques
(McLachlan & Liversidge 1970), South America and outlying islands, like-
wise in New Zealand (Dwight 1925) and also in south-western Madagascar
(Delacour 1932; Rand 1936; Milon 1948, 1950).
. The two species are very similar in appearance. Measurements of wing,
tail, tarsus and length of culmen per se are of little use in distinguishing them
as there is a wide range of overlap (Dwight 1925 ; McLachlan & Liversidge
1970). However the bill and tarsus are more slender in L. fuscus, culmen more
than three times depth at the base, while in L. dominicanus the culmen is less
than three times the depth at the base (Dwight 1925 : 126). This also applies
to the depth at the angle of the lower mandible and is a useful field character.
The yellow legs and feet of L. fuscus compared with the greenish-yellow or
bluish-grey legs and feet of L. dominicanus is probably the most diagnostic
field character. In adult birds the black mantle of L. dominicanus is similar to
that of L. fuscus in the field. Birds with slate-grey or paler mantles could not
63
be confused with L. dominicanus as these would either belong to the British
race L. f graellsii or races from northern Europe or Eurasia. However,
workers who are familiar with the heavier build and waddling gait of L.
dominicanus of coastal South Africa would have little trouble in distinguishing
the more slender L.fuscus in the field.
Recent observations show that L.fuscus occurs south of the Zambesi more
frequently than previously realised and that individuals have been recorded
near Durban at 30 S (Clancey in lift.). Several records of black-backed gulls
from inland waters of South Africa have been attributed to L. dominicanus
and these identifications were rightly questioned by Harwin (1952). Thus
there has been some confusion in the past concerning the field characters of
L. fuscus and L. dominicanus especially toward the southern edge of the range
of the former in Africa. In the light of the data gathered in Tables 1 and 2 it
is more likely that the gulls previously reported as L. dominicanus (Table 3)
TABLE 1
Larus fuscus: Collected material from southern and central Africa
Date
Plumage
Locality
Museum in which housed
Zambia
19.9.59
imm.*
Kabeti
National Museums of Rhodesia,
Bulawayo.
29.12.69
imm.
Lochinvar
National Museum of Zambia,
Livingstone.
Caprivi Strip
n. 12. 61
imm.
Lake Liambesi
National Museums of Rhodesia,
Bulawayo.
Rhodesia
14.10.70
imm.
Lake Kariba
National Museums of Rhodesia,
Bulawayo.
South Africa
28.12.53
imm.
Bon Accord Dam,
Pretoria.
Transvaal Museum, Pretoria.
* moulting into adult plumage.
from far inland in South Africa (Joubert 1943; Prozeski & Campbell 195 1;
Harwin 1952) were in fact L.fuscus. Indeed both editions of Roberts birds of
South Africa (McLachlan & Liversidge 1957, 1970) erroneously report a
bird collected at Bon Accord Dam, Pretoria as L. dominicanus. Dr. A. C. Kemp
of the Transvaal Museum, who examined the specimen at my request,
reported (in litt.) that "the bird is, by comparison, certainly Larus fuscus,
with markedly less heavy bill and smaller size."
The situation regarding the identification of black-backed gulls at Beira,
Mocambique, has been equally confusing. Bolster (1933) and Moreau (1938)
both record seeing L. dominicanus at this locality (Table 3). Winterbottom
(1936) recorded black-backed gulls at Beira but stated that "I cannot pretend
to be competent to differentiate between the two species in the field." Benson
(1948) saw several gulls which "were either L. dominicanus Lichtenstein or a
race of L.fuscus Linnaeus." Worth (i960) recorded several black-backed gulls
at Beira harbour during July and August 1959. These were identified as L.
dominicanus mainly on the grounds that "one would not expect these northern
migrants (L.fuscus) in southern waters in July and August." Since L.fuscus
has now been recorded from southern Africa virtually throughout the year
(Tables 1 and 2) this latter statement does not apply. Clancey (1969) points
out that L. dominicanus has been recorded as far north as Lourenco Marques
64
TABLE 2. Visual observations assessed as Larus fuscus
Date
No.
Plumage
Locality
Source
Malawi
Feb.
—
—
Linthipe, Lake Malawi
Benson (1953)
Nov-Mar
"Not uncommon'
' —
Lake Malawi
>> ■>■>
3.12.60
1
ad.
Lake Chilwa
Long (1962)
21.12.60
4 1
—
>> •>■>
21.10.59
I
imm.
Fort Johnston, Shire River „ „
14. 10.61
"Few"
—
Kota Kota, Lake Malawi
>> >>
May i960
4
—
„
Zambia
Jan-Mar
—
—
Libonda and between
Silonga and Lukulu,
Zambesi River
White &
Winterbottom
(1949)
—
1
—
Itawa Swamp
23.7.65
1
ad.
Mfuwe, Luangwa River
Benson et a/.
(1970)
16.9.64
1
imm.
Lochinvar
R. J. Dowsett
(in litt.)
13.3.72
2
i
ads.
imm.
Sumbu, Lake Tanganyika
>> >>
[Benson et al. (1971, The birds of Zambia. London: Collins) summarised the local records,
and included a few additional to those above — Ed.]
Botswana
22.9.70 1
Mozambique
12.3.72 1
Rhodesia
19.10.52 1
3.4.60 i 2
5.6.68 1
9.6-31.7.67 1
4.10.68 1
22.12.69 1
„ 1
16.12.70 i 2
20.1. 71 1
16.9.71 1
10. 12. 71 3
July 1973 1
21.5.66 1
8.3.70 1
April 1970 1
13. 2.71 1
Nov7o-May7i 1
Jan-May 71 1
28.1.72 1
3
13.7.72 1
South Afrka
Dec 72-Jan 73 1
1972 r
ad.
Lake Ngami
A. J. Tree
(in litt.)
ad.
Beira
B.G.D.
ad.
Pasi Dam, Gatooma
R. K. Brooke
(in litt.)
imm.
Chirundu
Tree (1961)
ad.
Chikwenya Is.,
G. R. Thomson
Zambesi/Sapi junction
(in litt.)
imm.
Lake Kariba
B.G.D.
ad.
,, ,,
>»
ad.
>> >>
imm.
„ ,,
>>
imm.
,, ,,
»
imm.
»» >j
G. W. Begg
(in litt.)
imm.
,, ,,
,, ,,
ads.
,, ,,
,, ,,
imm.
n ,,
tt
ad.
»
A. J. Conway
(in litt.)
imm.
Lake Mcllwaine,
R. K. Brooke
Salisbury
(in litt.)
imm.
,, ,,
» >>
ad.
„
R. A. Bourlay
(in litt.)
ad.
>> >>
R. I. G. Attwell
(in litt.)
ad.
>> >>
B. D. Connock
(in litt.)
imm.
,, „
>> >j
ad.
5> 5>
R. K. Brooke
(in litt.)
imm.
>> J>
>> ,,
ad.
Tree (1973)
ad.
Umgeni River Mouth,
P. A. Clancey
Durban
(in litt.)
ad.
Natal Coast
,, ,,
1 "moulting" (Long 1962) 2 moulting into adult plumage
3 A live crippled bird in the aviaries, Mitchell Park, Durban.
TABLE 3
Black-backed gulls of uncertain identity or identified as Larus dominicanus now considered
more likely to have been L. fuscus
Date
No.
Plumage Locality
Source
Mozambique
24.3.32
8-9.3.35
"several"
"several"
— Beira
ad. and imm. „
Bolster (1933)
Winterbottom (1936)
29.11-2.12.35
"common"
>> »
17-18. 5. 37
28.1.47
July, Aug 59
South Africa
May 1 941
2-9
"several"
"several"
1
>>
ads.
ad. and imm. „
— Alexander Dam,
Geduld
Barberspan
>>
ad. Odendaalsrus
Moreau (1938)
Benson (1948)
Worth (i960)
Joubert (1943)
Feb 1952
24.1.59
1
1
Prozeski &
Campbell (195 1)
Harwin (1952)
Brooke (1960)
on the east coast but that "some of the black-backed gulls occurring at Beira
may be applicable to Larus fuscus Linnaeus rather than L. dominicanus. No
specimens exist at the present time in order to investigate this matter."
Clancey's caution is noted since an occasional movement of L. dominicanus
to as far north as Beira is not impossible but which I feel would be highly
unlikely. In addition the lack of confidence in the identifications and the
growing list of Z.. fuscus data from inland and coastal waters suggest that the
above records were very likely to have been L. fuscus (Table 3).
The occurrence of L. dominicanus in south-western Madagascar at between
24 and 2 5 ° S is important in discussion relating to movements on the east
coast of Africa. McLachlan & Liversidge (1970) state that it is a migrant from
the Cape to Natal and northwards. Rand (1936) gives the earliest date of
occurrence for Madagascar (in the extreme south-west) as 12 February and
the latest as 4 March 1930. This would also imply that it is a migrant and if
so South Africa would be the obvious origin of the birds. However, Milon
(1950) records seeing 45 birds on 26 January, also in the extreme south-west,
and the same author (1948) reports June and July observations on L. domini-
canus from this same area. A female collected on 19 July had the ovary
measuring 27X7 mm and the three bigger eggs measured 4-8-5-3 mm.
Further north in western Madagascar, despite years of experience, Father O.
Appert (in litt. to Benson) has only once seen a gull with a black mantle,
which he believes was L. dominicanus, at Lac Iotry (22 S) on 12 May 1964.
The presence of birds during summer and winter months together with the
suggestion of local breeding points to a resident population of/., dominicanus
in south-western Madagascar and not a migration from Africa.
On 1 2 March 1972, a single adult L. fuscus was seen together with several L.
cirrocephalus and a single L. leucophthalmus in the fishing harbour at Beira
(B.G.D.). This bird was flying above the other gulls and attempted to rob
them of food picked up from the water. In striking contrast to the sooty
black mantle and wings of L. dominicanus and the adult examples of L. f
fuscus personally seen on Lake Kariba this bird had a fresh slate-grey plumage
characteristic of the British race L.f. graellsii. This race of JL. fuscus generally
winters on the Atlantic coast of Africa and has not been recorded from the
inland waters of the continent (Archer & Godman 1937; Vaurie 1965;
Moreau 1972). However, the bird collected at Kabeti (Table 1) was identified
66
as "probably a pale individual of L. f fuscus, though showing some inter-
gradation with L. f graelsii" (Benson i960). It would be surprising that an
example of a western Palaearctic population should be found on the east
coast of Africa. While the Beira bird could perhaps have followed the coast
from the Mediterranean, Red Sea and southwards from Somaliland, the more
likely possibility arises that it belonged to part of the L. fuscus complex of
races from northern Europe or Eurasia. Indeed Voous (1965) gained the
impression that birds observed at Mombasa, Kenya, were intermediate
between L. f fuscus and L. f graellsii and suggested that they "probably
belonged to some of the dark-mantled populations of Larus fuscus antelius
from North Russia or Northwest Siberia." He mentioned too that they "did
not clearly show the characters of the lighter and slightly more eastern
populations more properly known as L. f heuglini" although one bird
"probably was slightly lighter."
Harris (1962) states that most of the British populations oiL. fuscus move
south between August and October and return during March and April.
However, 83% of British ringed birds recovered from the Atlantic coast of
Africa were in their first year and all June and July birds recovered abroad
were under breeding age. This trend must also apply to other western and
central European populations. It would seem, therefore, that the European
visitors to Africa arrive largely in immature plumage and probably remain
south until they approach maturity. While the bulk of the observations have
been made during the months October to March both adults and immature
birds have been recorded in southern and central Africa virtually throughout
the year (Tables 1 and 2). Harris (1970) points out "that some gulls do not
attain full adult plumage even when seven years old", thus the actual duration
of stay in Africa by individuals is uncertain.
Acknowledgements : I am very grateful to Mr. M. P. Stuart Irwin of the
National Museum, Bulawayo, and Mr. R. K. Brooke of the Durban Museum
and Art Gallery for reading and commenting on an earlier draft of the text.
I would also like to thank Dr. A. C. Kemp for examining material in the
Transvaal Museum, Pretoria, and my colleague Mr. G. W. Begg of the Lake
Kariba Fisheries Research Institute, Rhodesia, for collecting a specimen and
providing several sight records at my request. I am particularly indebted to
Mr. C. W. Benson for his criticism and contribution to the data on Mada-
gascar. My thanks are also due to Mr. P. A. Clancey, Durban Museum and
Art Gallery, and Mr. R. J. Dowsett, National Museum of Zambia, Living-
stone, for providing much supporting data.
References :
Archer, G. & Godman, E. M. 1937. The birds of British Somaliland and the Gulf of Aden 2.
London: Gurney & Jackson.
Benson, C. W. 1948. Notes from a sea voyage: December, 1946 — January, 1947. Ostrich 19:
150-151.
— 1953. A check list of the birds of Nyasaland. Blantyre: Nyasaland Society.
— i960. Recent records from north-western Northern Rhodesia. Bull. Brit. Orn. CI. 80:
106-112.
Benson, C. W., Brooke, R. K., Dowsett, R. J. & Irwin, M. P. S. 1970. Notes on the birds of
Zambia: Part V. Arnoldia {Rhodesia) 4: 1-59.
Bolster, R. C. 1933. Sea birds observed on an east coast voyage. Ostrich 4: 15-20.
Brooke, R. K. 1960. Waterfowl on the Orange Free State Goldfields. Ostrich 31: 27-29.
Clancey, P. A. 1969. A handlist of the birds of southern Mozambique. Part 1. Mems. Inst.
Invest, cient. Mocamb. 10, A: 145-303.
Delacour, J. 1932. Les oiseaux de la Mission Zoologique Franco-Anglo-Am ericaine a
Madagascar. Oiseau 2: 1-96.
D wight, J. 1925. The gulls (Laridae) of the world. Bull. Amer. Mus. Nat. Hist. 72: 63-401.
67
Harris, M. P. 1962. Migration of the British Lesser Black-backed Gull as shown by ringing
data. Bird Study 9 : 1 74-1 82.
— 1970. Abnormal migration and hybridization oi Larus argentatus and L.fuscus after inter-
species fostering experiments. Ibis 112: 488-498.
Harwin, R. M. 1952. Does the European Lesser Black-backed Gull migrate to South
Africa? Ostrich 23: 123-124.
Joubert, H. J. 1943. Some notes on birds. Ostrich 14: 1-7.
Long, R. C. 1962. The Lesser Black-backed Gull Larus fuscus in Nyasaland. Ostrich 33(3):
52 (as amended Ostrich 34: 52).
McLachlan, G. R. & Liversidge, R. 1957. Roberts birds of South Africa. Cape Town: S. Afr.
Bird Book Fund.
— 1970. Roberts birds of South Africa. Cape Town: John Voelcker Bird Book Fund.
Milon, Ph. 1948. Notes d'observation a Madagascar. Alauda 16: 55-74.
— 1950. Deux jours au Lac Tsimanampetsotsa. Nat. Malgache 2: 61-67.
Moreau, R. E. 1972. The Palaearctic- African bird migration systems. London & New York:
Academic Press.
Moreau, W. M. 1938. A transect census from London to Tanga via the Cape. Ostrich 9:
35-46.
Prozeski, O. P. M. & Campbell, T. 195 1. A day at Barberspan. Ostrich 22: 192-194.
Rand, A. L. 1936. The distribution and habits of Madagascar birds. Bull. Amer. Mus. Nat.
Hist. 72: 143-499.
Tree, A. J. 1961. Observations on migrating movements of some migratory birds in
Northern Rhodesia. Ostrich 32: 86-89.
— 1973. Lesser Black-backed Gulls on Lake Mcllwaine. Honeyguide 73 : 32.
Vaurie, C. 1965. The birds of the palearctic fauna. Non-passerines. London: H. F. & G. Wither-
by.
Voous, K. H. 1965. Summer records of Larus fuscus in Mombasa, Kenya. Ardea 53: 83.
White, C. M. N. & Winterbottom, J. M. 1949. Check list of the birds of Northern Rhodesia.
Lusaka: Govt. Printer.
Winterbottom, J. M. 1936. Notes on birds around Beira in 1935. Ostrich 7: 32-38.
Worth, C. B. i960. Notes on sea birds in harbours of Portuguese East Africa. Ostrich 31 :
i73- I 74-
Observations on a relationship between
Crested Guineafowl and Vervet Monkeys
by Geoffrey Hill
Received 1st February, 1974
The Vervet Monkey Cercopithecus aethiops is the object of a continuing field
study in a coastal forest on the Natal North Coast, a forest which is also the
southernmost station of the Crested Guineafowl Guttera edouardi. In this
forest there are four fairly large (ca. 1 5 individuals in each) groups of Vervet
Monkeys. When they feed in trees they are the highest mammals in the forest
and thus have the widest view of their surroundings. It appears that other
animals take advantage of this factor, as on many occasions Bushbuck
Tragelaphus scriplus, Blue Duiker Cephalophus monticolus^ Red Forest Duiker
C. natalensis, Common Duiker Sylvicapra grimmia, Cane Rats Thryonomys
swinderianus and Crested Guineafowl have been noted feeding directly below
the monkeys.
Crested Guineafowl, especially, are found below them more often than not.
They associate chiefly in the late afternoon at which time both animals have
their final feed before retiring to sleep. This association is advantageous to
the Crested Guineafowl as the Vervet Monkeys are very messy feeders and
are continually dropping food items and knocking down seeds and berries.
Thus, by standing under the feeding monkeys, the guineafowl receive a
constant shower of food, since many of the kinds of berries that the monkeys
eat are also eaten by the guineafowl. The monkeys' faeces also contain a
large quantity of undigested seeds which the Crested Guineafowl probably
eat (e.g. Lanlana camara, Melia a^ederach).
68
Bordering the forest is a sugar cane field in which the Vervet Monkeys
spend considerable time, breaking canes which they carry off to the trees at
the edge of the forest to eat. On most occasions the Crested Guineafowl
accompany the monkeys to the forest edge and then wander along in the
cleared fire-break which separates the forest from the sugar cane, where they
dig up grubs and bulbs for food. They seldom stray far from where the
monkeys are.
When they are beneath the monkeys, the Crested Guineafowl seem to be
less afraid of those things that would normally disturb them. While sitting
on the ground watching the Vervet Monkeys, I have had Crested Guineafowl
come within a metre or so of me, and on occasion they have approached my
dog closely. Even when I move, the guineafowl do not appear to be unduly
disturbed.
It is not only the guineafowl who derive advantages from associating with
the Vervet Monkeys : there is also a reciprocal advantage, since the monkeys
react to the alarm calls of the guineafowl. When a flock of Crested Guinea-
fowl are disturbed by my walking through the forest, they give a rapid
"clicking" call and, often, within a few seconds, the Vervet Monkeys bark
from some distance away. After 20 to 40 seconds, depending on how far away
they are, the monkeys appear as if to investigate what had disturbed the
guineafowl.
When I take a companion into the forest with me, the Vervet Monkeys
begin to bark as soon as they see him and normally the Crested Guineafowl
"click" excitedly and move away from the area as if they recognize that the
monkeys are barking at something which is a potential danger. But, on other
occasions, their curiosity appears to overcome them and they venture closer
to inspect the intruder. This foraging and warning relationship between the
Crested Guineafowl and the Vervet Monkeys appears to be unreported in
the literature.
I am obliged to Mr. R. K. Brooke for criticizing a draft of this communi-
cation.
A new subspecies of Accipiter tachiro
by M. Desjayes
Received 2$rd November, iyj}
In April and May 1971, the author made a collection of birds in the province
of Kaffa in southwestern Ethiopia. The region extending from western
Kaffa to Gore in the Illubabor province is covered with very dense unbroken
humid forest which appears to be the largest expanse of rain-forest east of the
Congo from which it is isolated by over 1000 km of dry savanna. As should
be expected, a certain number of species collected are darker than those from
the rest of Ethiopia, which is generally drier.
Of the better differentiated birds are two specimens, a male and a female, of
Accipiter tachiro. Both are notably darker below than any other specimens of
the species. The barring of the female is definitely brown, not rufous. The
flanks and thighs are also darker in the same proportion. The male is rufous
below but also darker. Both sexes are separable at a glance from specimens
of unduliventer (type locality Simien) from Ankobar, Shoa, Ali-Beret, Arussi
and Manaco (between Anadi and Alelu, southern Ethiopia), and differ even
more from unduliventer than do many specimens oisparsimfasciatus (type locality
Zanzibar). The former is described as "darker, with heavier barring below
69
more washed with chestnut" (Brown & Amadon 1968); this character may
show in a fair series but is not evident in a small collection and individuals
are not always separable, as also noted by White (1965). The two Kaffa speci-
mens are similar in size to unduliventer for which Brown & Amadon give:
wing o* 184, $ 216 mm, and thus smaller than sparsimfasciatus. Weight
appears to be a usually good indicative of size variations although this charac-
ter is little used in subspecinc taxonomy, in part because of the paucity of
data, although see Amadon (1943). The weights of two females, 408 and 509
g, given by Brown & Amadon (1968) are a good measure of the size of
sparsimfasciatus although they seem to be among the heaviest. These figures
must have been derived from Moreau (1944). Verheyen (1953) gives £ 230 g
for sparsimfasciatus from southern Zaire and Britton (1970) an identical
figure from Tanzania, also in the distribution range of sparsimfasciatus. These
weights as well as those from Kaffa specimens given below show the re-
markable size dimorphism in this species. For the dark population of south-
eastern Ethiopia, I propose the name:
Accipiter tachiro croi^ati, subsp. nov.
in honour of Dr. Leon Croizat (botanist, biogeographer and scholar, Member
of the Academy of Sciences of Caracas, Corresponding Member of the
American Museum of Natural History) who has devoted to ornithology a
sizeable portion of his important work.
Type: Adult male with testes enlarged. Locality: Afallo, altitude ca. 2050 m,
Ghera region, 36°2o' E, 7°45' N, Kaffa province, 1 May 1971. U.S.Nat. Mus.
No. 522426.
Paratype: Adult female with ovary developing. Locality: Challa, altitude
ca. 2100 m, about 20 km north of Afallo, 8 May. US. Nat. Mus. No. 522427.
Diagnosis: Differs from unduliventer and from all other subspecies of
Accipiter tachiro in being darker both above and below ("Verona brown" vs.
"pecan brown" of Ridgway's Color Chart, 191 2). Size smaller than sparsim-
fasciatus, equal to unduliventer. Tail spots large and distinct. Under tail-coverts
barred with grey in q*, partly and very lightly so in $.
Colours of bare parts: Male: Iris orange-yellow; orbital skin same but paler.
Bill: upper mandible blackish, lower dark brown at tip, bluish at base; cere
and commissures lemon-yellow. Legs and feet yellow ; soles paler and brighter
yellow; claws dark brown. Female: Iris and orbital skin yellow. Bill: upper
mandible blackish-slate, bluish at base of sides; cere and commissures
greenish yellow; lower mandible blackish-slate, bluish in the middle, pale
yellow at base. Legs, feet, soles and claws as in male.
Measurements: Wing $ 172, $ 217; tail <$ 150, $ 190; bill from skull <$ zz,
$ 25 ; tarsus £ 54, $ 60 mm. Weight 6* 145, ? 260 g.
Material examined: Specimens of unduliventer, sparsimfasciatus and nominate
tachiro in the U.S. National Museum and the American Museum of Natural
History.
Distribution: The rain-forest of southwestern Ethiopia.
Acknowledgments
I gratefully acknowledge the support of the National Geographic Society
who made the collecting possible. My thanks also go to Dr. Dean Amadon
who extended to me the use of the facilities of the American Museum of
Natural History, and has commented on this paper.
70
References :
Amadon, D. 1943. Bird weights as an aid in taxonomy. Wilson Bull. 55 : 164-177.
Britton, P. L. 1970. Some non-passerine bird weights from East Africa. Bull. Brit. Orn. CI.
90: 142-144.
Brown, L. & Amadon, D. 1968. Eagles, hawks and falcons of the world. London : Country Life
Books.
Moreau, R. E. 1944. Some weights of African and of wintering Palaearctic birds. Ibis 86:
16-29.
Verheyen, R. 1953. Exploration du Pare National de PUpemba. Fasc. 19. Oiseaux. Inst. Pares
Nat. Congo Beige, Bruxelles.
White, C. M. N. 1965. A revised check-list of African non-passerine birds. Lusaka: Government
Printer.
The eggs of the Golden Conure Aratinga guarouba
by Michael P. Walters
Received 4th January, 1974
There appear to be no references to the eggs of the Golden Conure in
literature beyond the statement by Snethlage (1935) that an egg removed
from the oviduct of a specimen collected on the Tocantins river on 13 th
October 191 2, was "almost round". This has been quoted in Meise (1963-4)
and by Forshaw (1973), without further comment. The collection of the
British Museum (Natural History) contains seven eggs of this species, a c/6
laid in August 1938 (reg. no. 1938.8. 1. 1-6) from the collection of the
Marquis of Tavistock, and a single egg recently acquired from the collection
of the late Col. F. E. W. Venning (reg. no. 1970.6.520). The Tavistock eggs
are roundish ovals, two of them being narrowly pointed at the small end.
They are smooth, but not glossy, and measure 35-4 x 25-7; 34-3 x 25-1;
32-9 x 26-5; 35-1 x 25-8; 32-2 x 25-2; and 32-7 x 26-5 mm. The
Venning egg is slightly smaller, 30-2 x 25-2 mm, and is almost round.
References :
Forshaw, J. M. 1973. Parrots of the world. Melbourne: Lansdowne Press.
Meise, W. (ed.) 1964. Schonwetter: Handbuch der Oologie. Berlin: Akademie Verlag. 8-9.
Snethlage, E. 1935. Beitrage zur Fortpfianzungsbiologie brasilianischer Vogel. Journ.
Ornith. 83: 532-562.
Reduced natal down in some emberizine species
by C. J. 0. Harrison
Received 1 8 th January, 1974
In describing the natal pterylosis of the Swallow-tanager Tersina viridis,
Collins (1973) has compared the extremely scanty down of this species with
the more plentiful down of other Thraupinae species and suggested that the
reduction of neossoptiles in that species and in the Violet Euphonia Tanagra
violacea is correlated with a nest in a cavity or closed site as opposed to the
more typical cup nest in an open site. He comments that further information
is needed on this feature of tanagers and allied species.
Recently observations were made on the captive breeding of Saffron
Finches Sicalis flaveola (Harrison 1973). This species normally nests in a cavity
or in the closed nest of a larger bird, and within the site builds a cup-type
nest. The site used by the captive birds was a small hollow log, and it was
only possible to examine one nestling at about six days old. This was almost
naked, but close examination showed a few tiny, sparse down tufts on the
middle of the dorsal tract, a few more on the upper wings, and one or two
over each eye.
Skutch (1954) cites the Yellow-faced Grassquit Tiaris olivacea as exceptional
in having completely naked nestlings, and the species builds a thick-walled,
domed nest with a small lateral entrance.
Both Sicalis and Tiaris belong to the Emberizinae, and the present evidence
suggests that in both subfamilies of the Emberizidae there is a reduction in
the natal down of species nesting in closed nests, or nests within cavities.
References :
Collins, C. T. 1973. The natal pterylosis of the Swallow-tanager. Bull. Brit. Orn. CI. 93:
I55-I57-
Harrison, C. J. O. 1973. Observations on the behaviour and breeding of the Saffron Finch,
Sicalis flaveola. Avicult. Mag. 79: 207-213.
Skutch, A. F. 1954. Life histories of Central American Birds. Pacif. Cst Avifauna 31 : 121.
Notes on the Lammergeyer Gypaetus barbatus
on Kilimanjaro
by I. H. Dillingham
Received 1st February, 1974
The summary of records of the Lammergeyer in East Africa by Schiiz (1973,
Bull. Brit. Orn. CI. 93 : 1 61-163) ^ as prompted me to publish two records of
probably the same individual of this species seen by myself on 23 rd and 26th
October 1956 at Engare Nairobi on the western slopes of Kilimanjaro at an
altitude of approximately 4500 feet.
On the first occasion I had a clear view of an adult Lammergeyer about 75
yards from me at a height of about 100 feet circling over farmland. It then
flew to a farm about half a mile away where it repeatedly circled low down
over some outbuildings before disappearing eastwards. On the second
occasion I was in the same general area when an adult was seen gliding
overhead, again at a height of no more than 100 feet, finally disappearing in a
northerly direction without circling.
These two records at Engare Nairobi would appear to be the lowest
altitude at which this species has been recorded on Kilimanjaro. This area is
largely devoted to cattle farming and is relatively flat. Despite frequent visits
to Engare Nairobi and the surrounding country during the period June 1956
to August 1958 this species was never encountered on any other occasion.
It may be noteworthy that all the records of the Lammergeyer on Kili-
manjaro mentioned by Schiiz (1958, Journ. Ornith. 99: 394-398 and 1973, op.
cit.) for which a date is quoted, plus my own records, fall within the period
late August to December. The Mawenzi summit of Kilimanjaro is the most
likely breeding area yet it is the most inaccessible and infrequently visited.
The preponderance of records in the second half of the year, nearly all of
which are on the west or south-west of the mountain, may represent post
breeding dispersal from the eastern summit Mawenzi. An alternative ex-
planation for this distribution of the Lammergeyer is that it breeds on Mount
Meru which is a mere 3 5 miles west to south-west from the Shira Plateau.
This mountain would appear to provide a much more suitable breeding
habitat for this bird with its numerous crags. There is evidence from King
(1973, Bull. Brit. Orn. CI. 93: 68) that the Lammergeyer occurs there, but
again it is a September record.
I am most grateful to Mr. C. W. Benson for assistance in the preparation
of these notes.
72
A discography of bird sound from the
Neotropical Region
by Jeffery Bos wall <& W. P. Freeman
Received jth December, 197}
This paper is the sixth to be published of a number of projected papers, each
covering a major zoogeographical region of the world. For the Palearctic see
Boswall (1964, 1966, 1969a, 1969b, 1970, 1 971) and Sellar (1973); for
Australasia see Boswall (1965); for the Ethiopian Region Boswall & North
(1967); for the Antarctic Boswall & Prytherch (1969); and for the Oriental
Region Boswall (1973). A Nearctic discography is in final draft (Boswall &
Kettle in press).
The earliest Neotropical recordings of birds were probably those made on
Barro Colorado Island, Panama, during World War II (Wyring, Allen &
Kellogg 1945; Kellogg & Allen 1950, no. 1 in the following discography;
and Asch & Ramsey 1952, no. 2 in the following discography).
Since that time no doubt many workers have recorded birds in the Neo-
tropical region. To try to bring together details of their results would not be
an easy task. The primary purpose of this paper is to give details of published
recordings; however some indication of the unpublished recordings that have
been made is not entirely out of place.
One of the outstanding contributions is that of the indefatigable L. Irby
Davis who has recorded well over 350 species in Mexico, many of which are
held at the Balcones Research Centre, University of Texas, Austin. Burton
(1969), the BBC's wildlife sound librarian, includes reference in his main
catalogue to a Least Bittern Ixobrychus exilis recorded in Colombia, and a
Banded Wren Tbryothorus pleurost ictus recorded in Mexico. In the first supple-
ment to the main catalogue the same compiler (Burton 1971) refers to a range
of recordings by Ian Strange from the Falkland Islands in the South Atlantic,
as well as to some Trinidad recordings by D. W. Snow, and to one or two
from Little Tobago (also in the West Indies) by Robina Gyle-Thompson.
Burton's second supplement (1972) refers to a few recordings made in
Jamaica by Oxford Scientific Films Ltd., to some additional Falkland Island
recordings by Ian Strange and Tony Morrison, and to recordings from the
Galapagos Islands in the Pacific made by Roger Perry and Bob Wade.
Further recordings from the Galapagos Islands and from Barro Colorado
Island in the Panama Canal have recently been incorporated into the BBC's
Sound Archives (Burton 1973). And it is hoped that recordings of about forty
bird species made in Argentina between October 1971 and March 1972 by
Robin J. Prytherch, a member of the BBC Wildlife Expedition to Southern
South America, may eventually be included in the BBC Sound Archives.
Recordings of ca. 20 species made in Buenos Aires Province of Argentina
by Donaldo Maclver Junior have been deposited with the British Library of
Wildlife Sounds at the British Institute of Recorded Sound in London.
A dozen or so recordings made in Brazil are preserved in the Library of
Natural Sounds at the Laboratory of Ornithology at Ithaca, New York State,
U.S.A., as are many Caribbean recordings of George B. Reynard, and no
doubt many others from the region in question.
The discography
1. Kellogg, P. P. & Allen, A. A. 1950. Jungle Sounds. One 12-inch 78 r.p.m. disc no. BL
4219-20. Comstock Publishing Co., Inc., Ithaca, N.Y., U.S.A. [11 birds, 1 mammal and 7
amphibians, recorded in Panama.]
73
2. Asch, M. & Ramsey, F., Jr. 1952. Sounds of a Tropical Rain Forest in America. One 12-
inch 33 r.p.m. disc no. FX 6120. Produced for the American Museum of Natural History
by Folkways Records, 701, 7th Avenue, New York City, N.Y., 10036, U.S.A. [About 22
birds, 1 mammal, a few insects, and about 6 amphibians, recorded in Panama and at the
Bronx Zoo, New York City.]
3. Davis, L. Irby. 1958. Mexican Bird Songs. One 12-inch 33-3 r.p.m. disc. Laboratory of
Ornithology, Cornell University, Ithaca, N.Y., U.S.A. [74 birds.]
4. Gunn, W. W. H. 1959. A Day at F lores Morades, no. 5 in the Sounds of Nature series. One
12-inch 33-3 r.p.m. disc, no. T46558-9. Federation of Ontario Naturalists, Don Mills,
Ontario, Canada. [31 birds, 1 mammal and 2 insects. Revised and reissued as no. 35.]
5. Coelho, M. About i960. Hunting all through Brazil (Portuguese). One 12-inch 78 r.p.m.
disc, no. PR244. Gravacos Electricas, Sao Paulo, Brazil. [Instrumental imitations of about
12 birds. See Mitchell (1957: 32).]
6. Zuber, C. i960. Galapagos (French). One 7-inch 45 r.p.m. disc, no EX 253. From La
Boite a Musique, 133 Boulevard Raspail, Bam, Paris, France. [7 birds and 1 mammal.]
7. Frisch, J. D. 1961. Songs of the Birds of Brazil (Portuguese). One 12-inch 33-3 r.p.m. disc,
no. SCLP 10502. SOM, Sao Paulo, Brazil. [33 birds, 2 insects and 1 amphibian.]
8. Frisch, J. D. 1962. Songs of the Birds of Brazil (Portuguese). One 12-inch 78 r.p.m. disc,
no. S596. SOM, Sao Paulo, Brazil. [9 birds from the 33 species on no. 7.]
9. Frisch, J. D. 1962. Songs of Birds of South America (Spanish). One 12-inch 33-3 r.p.m.
disc, no. LP 040. Copacabana, for El Palacio de la Musica S.A., Venezuela. [34 birds; a
Spanish version, with changed title, of no. 7.]
10. Frisch, J. D. 1962. Songs of the Birds of Brazil (Italian). One 12-inch 33-3 r.p.m. disc, no.
Sabrina SA 10.001. Compagnia Discografica Sabrina, R. Mario Pagano 61, Milan, Italy.
[34 birds; an Italian version of no. 7].
11. Frisch, J. D. 1962. Songs of Birds of Brazil (Esperanto). One 12-inch 33-3 r.p.m. disc, no.
SCLP 10523. SOM, Sao Paulo, Brazil. [34 birds; an Esperanto version of no. 7.]
12. Frisch, J. D. 1962. Songs of Birds of South America (Spanish). One 12-inch 33-3 r.p.m.
disc, no. OL 7012. Opus, Surco Industrias Musicales S.R.L., Argentina. [34 birds; an
Argentinian version of no. 7.]
13. Frisch, J. D. 1962. Voices of the Brazilian Jungle (Portuguese). One 12-inch 33-3 r.p.m.
disc, no. SCLP 105 13, SOM, Sao Paulo, Brazil. [24 birds, 1 mammal, 2 insects and 1
amphibian.]
14. Frisch, J. D. 1963. Voices of the Amazon, with the Legendary Song of the Musician Wren
(Portuguese). One 12-inch 33 -3 r.p.m. disc, no. SCLP 10525. SOM, Sao Paulo, Brazil. [28
birds, 4 mammals, 1 insect and 2 amphibians.]
15. Frisch, J. D. 1963. Echoes of the Green Hell (Portuguese). One 12-inch 33-3 r.p.m. disc,
no. SCLP 10527. SOM, Sao Paulo, Brazil. [20 birds, 4 mammals, 2 reptiles, 1 amphibian, 4
fish, 6 insects and 1 arachnid.]
16. Schwartz, P. 1964. Bird Songs from the Tropics, Vol. 1 of Naturale^a Vene^plana. One 12-
inch 33-3 r.p.m. disc. Instituto Neotropical, Apartado 4640 Chacao, Caracas, Venezuela. [40
birds.]
17. Salibi, Dr. Jose, Jr. 1964. The Chestnut-bellied Rice Grosbeak (Portuguese). One 7-inch
33-3 r.p.m. disc, no. C.S. 70.071. Discos RGE Ltda, Rua Paula Souza 181, 2.0 e 5.0 andares,
Sao Paulo, Brazil. [One bird in captivity.]
18. Frisch, J. D. 1964. Songs of Birds of Brazil (Portuguese). One 7-inch 45 r.p.m. disc, no.
COP 73037. Copacabana, Discos de Radio Triunfo, Lda, Porto, Portugal. [7 birds; a
Portuguese edition of part of no. 7.]
19. Frisch, J. D. 1965. Symphony of the Hinterland (Portuguese). One 7-inch 33-3 r.p.m. disc,
no. 5001. SOM, Sao Paulo, Brazil. [Human musical arrangements intermingled with the
songs of unnamed Brazilian birds; of little ornithological importance.]
20. Frisch, J. D. 1965. Symphony of the Dawn (Portuguese). One 7-inch 33-3 r.p.m. disc, no.
5002. Sabia, SOM, Avenida Casper Libero, 58-12 Andar, Sao Paulo, Brazil. [Human
musical arrangements intermingled with the songs of unnamed Brazilian birds.]
21. Frisch, J. D. 1965. Symphonies of the Canaries (Portuguese). One 7-inch 33 -3 r.p.m. disc,
no. 50.004. SOM, Avenida Casper Libero, 58-12 Andar, Sao Paulo, Brazil. [One captive
bird with a background of organ music]
22. Frisch, J. D. 1965. Symphony of the Brazilian Birds (German). One 7-inch 33-3 r.p.m. disc,
no. 5004. Produced for Varig Airlines by SOM Ind. e Com. S.A., Avenida Casper Libero,
58-12 Andar, Sao Paulo, Brazil. [Human musical arrangements intermingled with songs of
unnamed Brazilian birds.]
23. Frisch, J. D. 1965. Symphony of the Brazilian Birds (Italian). One 7-inch 33-3 r.p.m. disc,
no. 5005. Produced for Varig Airlines by SOM Ind. e Com. S.A., Avenida Casper Libero,
58-1 2 Andar, Sao Paulo, Brazil. [Human musical arrangements intermingled with the songs
of unnamed Brazilian birds.]
74
24. Frisch, J. D. 1965. Symphony of the Brazilian Birds (French). One 7-inch 33 -3 r.p.m. disc,
no. 5006. Produced for Varig Airlines by SOM Ind. e Com. S.A., Avenida Casper Libero,
58-12 Andar, Sao Paulo, Brazil. [Human musical arrangements intermingled with the songs
of unnamed Brazilian birds. J
25. Frisch, J. D. 1965. Symphony of the Brazilian Birds (Portuguese). One 7-inch 33-3 r.p.m.
disc, no. 5007. Produced by Varig Airlines by SOM Ind. e Com. S.A., Avenida Casper
Libero, 58-12 Andar, Sao Paulo, Brazil. [Human musical arrangements intermingled with
songs of unnamed Brazilian birds.]
26. Frisch, J. D. 1965. Symphony of Brazilian Birds. One 7-inch 33-3 r.p.m. disc, no. 5008.
Produced for Varig Airlines by SOM Ind. e Com. S.A., Avenida Casper Libero, 58-12
Andar, Sao Paulo, Brazil. [Human musical arrangements intermingled with songs of
unnamed Brazilian Birds.]
27. Frisch, J. D. 1965. Symphony of Brazilian Birds (Japanese). One 7-inch 33-3 r.p.m. disc,
no. 5010. Produced for Varig Airlines by SOM Ind. e Com. S.A., Avenida Casper Libero,
58-12 Andar, Sao Paulo, Brazil. [Human musical arrangements intermingled with songs of
unnamed Brazilian birds.]
28. Frisch, J. D. 1965. The Legendary Paradise of the Indians (Portuguese). One 12-inch 33-3
r.p.m. disc, no. SCLP 10.534. SOM, Avenida Casper Libero, 58-12 Andar, Sao Paulo,
Brazil. [35 birds, 2 mammals, 1 insect and 1 amphibian.]
29. Frisch, J. D. 1965. Bird Songs of Brazil and Other Jungle I 'oices. One 7-inch 45 r.p.m. disc,
no. EP 4618. Produced for Varig Airlines by SOM, Ind. e Com. S.A., Avenida Casper
Libero, 58-12 Andar, Sao Paulo, Brazil. [14 birds, 1 fish, 1 insect and at least 1 amphibian.]
30. Schwartz, P. 1965. Sounds of the Llanos I. Vol. 2 of Naturale-^a Vene-^plana. One 12-inch
33-3 r.p.m. disc with descriptive booklet. Instituto Neotropical, Apartado 4640 Chacao,
Caracas, Venezuela. [68 birds, 5 mammals, 2 amphibians and 1 insect.]
31. Schwartz, P. 1965. Sounds of the Llanos II. Vol. 3 of Naturale~a I 'ene^plana. One 12-inch
33-3 r.p.m. disc with descriptive booklet. Instituto Neotropical, Apartado 4640 Chacao,
Caracas, Venezuela. [About 58 birds, including some 23 not recorded on Vols. 1 and 2.|
32. Frisch, J. D. 1966. Symphony of the Brazilian Birds (Portuguese). One 12-inch 33-3 r.p.m.
disc, no. SCLP 10.538. SOM, Ind. e Com. S.A., Avenida Casper Libero, 58-12 Andar, Sao
Paulo, Brazil. [Human musical arrangements intermingled with songs of unnamed Brazilian
birds, combining on one 12-inch disc the items appearing on the series of symphonies
produced in 1965 for Varig Airlines.]
33. Frisch, J. D. Probably about 1967. Symphony of the Birds. One 12-inch 33-3 r.p.m. stereo
disc, no. SE4442. MGM Records Division, Metro-Goldwyn-Meyer Inc., 1350 Avenue of
the Americas, New York City, U.S.A. [An English language version of no. 32.]
34. Frisch, J. D. Probably about 1967. Christmas Symphony. One 7-inch 33-3 r.p.m. disc,
no. Sabia Compacto 5014. SOM, Ind. e Com. S.A., Avenida Casper Libero, 58-12 Andar,
Sao Paulo, Brazil. [Human musical arrangements intermingled with the songs of unnamed
Brazilian birds.]
35. Gunn, W. W. H. 1968. A Day at Flores Morades. One 12-inch 33-3 r.p.m. disc, no. 5 in
Sounds of Nature series, and no. T46558-9. Federation of Ontario Naturalists, Don Mills,
Ontario, Canada. [31 birds, 1 mammal and 2 insects recorded in Venezuela. A revised and
corrected edition of no. 4.]
36. Reynard, G. B. 1969. Caribbean Bird Songs. One 12-inch 33 -3 r.p.m. disc. P. P. Kellogg,
Cornell Laboratory of Ornithology, Ithaca, N.Y., U.S.A. [54 birds recorded in Puerto Rico
and the Virgin Islands.]
37. Lindblad, Jan. 1970. In Green Paradise (Swedish). One 12-inch 33-3 r.p.m. disc, no.
STP LP 102. Bonniers, Sweden. [38 birds, 10 mammals and 1 insect recorded in Guyana.]
38. Roche, J. C. Probably 1970. The Bird as Musician. No. 7: Rufous-throated Solitaire and
Bare-eyed Thrush. One 7-inch 45 r.p.m. disc, no. P 039. L'Oiseau Musicien, 04-Aubenas-les-
Alpes, Haute-Provence, France. Obtainable also from Discourses Ltd., 34 High Street,
Royal Tunbridge Wells, Kent, U.K. [2 birds.]
39. Frisch, J. D. Probably about 1970. Birds, Jungle, Melody (Portuguese). One 12-inch 33-3
r.p.m. disc, no. SCLP 10543. SOM, Ind. e Com. S.A., Rua Franca Pinto, Sao Paulo, Brazil.
[Side 1 records unnamed species of the Amazonian forest, species of the central plateau
and of the southern forest, and one captive bird accompanied by human music. Side 2
consists of six human musical arrangements intermingled with songs of unnamed Brazilian
birds.]
40. Frisch, J. D. Probably about 1971. Sounds of the Trans- Amazonian Highway and Thrush
Songs (Portuguese). One 12-inch 33-3 r.p.m. stereo disc, SOM, Ind. e Com. S.A., Rua
Eugenia S. Vitale, 173, Sao Bernardo do Campo (Rudge Ramos), Sao Paulo, Brazil. [Side 1
consists of 4 bands: the first records dawn sounds in the Amazonian forest, the second
Indian songs and a flute, the third sounds of nightfall in the forest followed by reveille
with an army unit, and the fourth sounds and music representing the opening up of the
75
backwoods. On Side 2 the songs of five Brazilian thrushes are given at length, but the
species on Side 1 are not identified by name when they are heard.]
41. Roche, J. C. 1971. Birds of the West Indies, 1: The Lesser Antilles, from Grenada to Guade-
loupe (French and English). One 12-inch 33-3 r.p.m. disc, no. G.06. L'Oiseau Musicien,
04 Aubenas-les-Alpes, Haute-Provence, France. Also obtainable from Discourses Ltd.,
34 High Street, Royal Tunbridge Wells, Kent. U.K. [More than 13 birds; 4 insects and 3
amphibians.]
42. Gibson, D. and others. 1972. Cries of the Wild, authentic nature sounds from the Audubon
Wildlife Theatre Series. One 12-inch 33-3 r.p.m. disc, no. T-57169. K.E.G. Productions,
556 Church Street, Toronto, Canada. [Side B, Band 3, running 3 minutes 57 seconds, gives
5 birds and 1 mammal recorded in Venezuela by W. W. H. Gunn, and an Amazon "atmos-
phere" recording by L. KofHer.]
43. Coffey, Ben, B., Jr. and Evans, E. R. Probably 1972. Songs of Mexican Birds. One 3-25
inch diameter reel of 7 -5 i.p.s. tape. Ben B. Coffey, 672 N. Belvedere, Memphis, Tenn.
38107, U.S.A. [Over 24 birds, 1 mammal and 1 amphibian.]
References :
Boswall, Jeffery. 1964. A discography of Palearctic bird-sound recordings. Brit. Birds 57,
special supplement: 1-63.
— 1965. A catalogue of tape and gramophone records of Australasian Region bird-sound.
Emu 65 : 65-74.
— 1966. New Palearctic bird-sound recordings in 1964-65. Brit. Birds 59: 27-37.
— 1969a. New Palearctic bird-sound recordings during 1966-67. Brit. Birds 62: 49-65.
— 1969b. New Palearctic bird-sound recordings during 1968. Brit. Birds 62: 271-281.
— 1970. New Palearctic bird-sound recordings during 1969. Brit. Birds 63: 324-332.
— 1971. New Palearctic bird-sound recordings during 1970. Brit. Birds 64: 431-434.
— 1973. A discography of bird-sound from the Oriental Zoogeographical Region. Bull.
Br. Orn. CI. 93: 170-172.
Boswall, Jeffery & North, Myles, E. W. 1967. A discography of bird-sound from the
Ethiopian Zoogeographical Region. Ibis 109: 521-533.
Boswall, Jeffery & Prytherch, Robin, J. 1969. A discography of bird-sound from the
Antarctic. Polar Record 14(92): 603-612.
Boswall, Jeffery & Kettle, Ron. In press. A discography of bird-sound from the Nearctic
Region.
Burton, John. 1969. Catalogue of Natural History Recordings, BBC.
— 1 97 1. First Supplement to the 1969 Catalogue of Natural History Recordings, BBC.
— 1972. Second Supplement to the 1969 Catalogue of Natural History Recordings, BBC.
— 1973- Third Supplement to the 1969 Catalogue of Natural History Recordings, BBC.
Mitchell, Margaret, H. 1957. Observations on birds of south-eastern Brazil. University of
Toronto Press.
Sellar, Patrick. 1973. New Palearctic bird-sound recordings in 1971-72. Brit. Birds 66: 303-
310.
Wyring, C. F., Allen, A. A. & Kellogg, P. P. 1945. Jungle Acoustics. O.S.R.D. Reports 4699
and 4704, published privately from the Division of Acoustical Research, Rutgers
University, Brunswick, New Jersey, U.S.A.
Nearctic waders in Sierra Leone— Lesser Golden Plover
and Buff-breasted Sandpiper
by G. D. Field
Received 19th February, 1974
There are few records of American waders in tropical Africa but, judging by
the numbers reported annually from Europe, this must partly reflect lack
of ornithologists rather than birds. The only records for West Africa given
by Mackworth-Praed & Grant (1970) are single occurrences of Calidris
melanotos (Principe Island) and Tringa solitaria (Cabinda). Since then Tringa
flavipes has occurred at Lagos (Wallace 1969) and Numenius phaeopus hudsonicus
76
at Freetown (Sudbury & Field 1972). An example of Pluvialis dominica at
Lagos (Heigham 1969) was thought to be of the Siberian race, P. d.fulva.
On the nights of November 3rd and 4th 1973 exceptionally violent storms
hit Freetown, bringing in their wake the first real rush of land migrants,
particularly Sylviidae. It is possible that these storms were also responsible
for the appearance of some scarcer waders, Knot Calidris canutus, first seen
November 7th, and Oyster-catcher Haematopus osfralegus, first seen November
i 5 th.
On the evening of November 6th at high tide J. B. Smart and I noted a
small group of waders standing on the causeway along the edge of the
muddy, mangrove-margined Aberdeen Creek at the north-west tip of the
Freetown Peninsula. Most birds were Charadrius hiaticula and Tringa hypoleucos
with one Grey Plover Pluvialis squatarola and beside it a bird which on its own
might have been dismissed as P. squatarola but, seen alongside, obvious
differences in size and stance proclaimed it a 'golden' plover. We were able
to get to fifteen yards range and note plumage details, later flushing it so that
the under-wing could be observed. It was located again on November 7th,
nth and 12th, on the latter two occasions on the mud, extremely tame and
coming, as the tide rose, to within a few yards of me, associating here with
Redshanks Tringa totanus and Greenshanks Tringa nebularia, wading up to its
thighs in the water and eventually flying off solitarily.
Compared with the Grey Plover this bird was smaller and more delicate
in all particulars, especially bill, always standing erect instead of in the hun-
ched attitude typical of the Grey Plover, and in flight its wings were narrower
and relatively longer, giving a more elegant appearance. Salient points:
forehead, broad streak over eye, face and neck buff with a dusky patch on ear
coverts; upperparts streaked and spangled dark and pale (white rather than
gold); below buff with greyish feather centres on the breast, lower belly
white. In flight wing dark above with no bar, below at a distance it looked all
buff; closer, a distinction visible between the greyer axillaries and the palish
buff rest of wing. It appeared to be a first winter bird, corresponding closely
to the illustration of the American juvenile depicted on plate 121 (Witherby
et al. 1940) except that the forehead was buff and the underparts less heavily
marked.
On November nth after the plover had flown, while watching the waders
congregating on a still unflooded patch of mud, I noted a single bird fly in
and land among a group of Grey and Ringed Plovers. Almost immediately
the latter took off and flew out to sea, the strange wader accompanying them.
On the ground it was taller than the Ringed Plovers, though in the air it
seemed roughly the same size. All I had time to note was dark wings and tail
with no white, buff below, and yellow legs. Searching on November 12th
and 13th brought no sign of it, but on November 15 th a very dark wader
flew from the mangroves and landed on the beach among a group of Sander-
lings Calidris alba and an Oyster-catcher. Wary at first, it later ignored me
almost entirely and let me approach to within ten yards, running along the
tide edge and refusing to fly when I wanted to flush it. Later it flew to open
sandy grass beside the road and was still there when I left but was not seen
subsequently.
As this is the first record for Africa of a Buff-breasted Sandpiper Tryngites
subruficollis I give a full description : general appearance on the ground, a very
buff bird with clearly patterned dark and buff upperparts, short black bill and
yellow legs. It stood much higher than the Sanderlings, thus looking a bigger
77
bird, and appeared to be of the general size and height of a Wood Sandpiper
Tringa glareola. In detail : crown brown-streaked with a darkish streak down
centre of nape; whole of rest of head, forehead, supercilium, cheeks, neck,
plain buff, the impression being that the area immediately round the eye was
paler than the rest. Wings boldly patterned dark and buff, the scapulars
particularly variegated with dark feather centres and broad buff margins.
Whole underside pale buff, sides of breast spotted with grey-brown at close
view; under tail coverts paler. Eye dark, bill black, rather stout and appearing
slightly down-curved, a little longer than the length of the head. Legs rather
bright yellow. In flight from above, dark with no discernible wing-bar and
no white in the tail though this was patterned when spread on landing.
Underside of wing wholly white, in marked contrast with breast and upper-
side. Wings rather long and pointed. At rest the bird stood high with a well-
developed neck. It both walked and ran, feeding at the tide's edge by turning
over small pieces of stranded seaweed, picking them up in the beak and
shaking them, often dropping them and giving an exaggerated jump back-
wards as though in alarm.
Without access to skins I cannot be categorical about the race of P.
dominica, but its appearance in conjunction with an undoubted American
species makes it reasonable to suggest that it too was of American origin,
the race P. d. dominka.
References:
Heigham, J. B. 1969. Birds observed in Victoria Island, Lagos. Bull. Orn. Soc. Nigeria 6
(No. 24): 109-119.
Mackworth-Praed, C. W. & Grant, C. H. B. 1970. African handbook of birds. Series III, Vol.
1. London: Longmans.
Sudbury, A. & Field, G. 1972. An African record of the Hudsonian Curlew Numenius
phaeopus hudsonicus Latham. Bull. Brit. Orn. CI. 92: 148-9.
Wallace, D. I. M. 1969. Lesser Yellowlegs at Lagos: a species new to Nigeria. Bull. Orn.
Soc. Nigeria 6 (No. 22): 58.
Witherby, H. F., et al. 1940. The handbook of British birds 4. London: Witherby.
The Little Egret Egretta gar^etta in Wallacea
by C. M. N. White
Received 26th January, 1974
Little Egrets have been recorded from at least fifteen islands in Wallacea*
mainly from Celebes and some adjacent islands and from north and south
Moluccas. Records from the Lesser Sunda and South West islands are
scantier. Amadon & Woolfenden (1952: 12-13) refer these records to E. g.
nigripes with legs and feet concolorous and black, which occurs from Java
and Borneo to Australia and New Guinea.
In recent years it has become apparent from ringing recoveries that the
nominate E. g. gar^etta with yellow feet, breeding in Japan, winters in
numbers in the Philippine Islands. This indicated that a new examination of
birds from Wallacea was desirable to ascertain whether any yellow footed
examples, presumptive migrant gar^etta, occur there. I am grateful to Dr.
K. C. Parkes for information about his study of Philippine birds which has
subsequently (1973: 14-15) been published. In addition to yellow footed
gar^etta, he found evidence of a breeding population in the Philippines with
78
greenish yellow feet spotted with black. This he suspected might represent
a third form, perhaps undescribed. Parkes had seen few breeding specimens
from the Philippines, but found that Tweeddale and McGregor had given
descriptions of such birds with spotted feet which were consistent with the
few specimens he had examined.
I am also grateful to Dr. G. F. Mees for information and comments about
specimens in the Leiden Museum. Eleven recently collected and undoubtedly
breeding birds from Formosa all have black feet as in nigripes. It is thus
strange that no black footed breeding birds have been found in the Philip-
pines, and there would seem to be need for more investigation of the foot
colour of Little Egrets breeding there. This is indicated by the situation in
Wallacea.
Most specimens in Leiden are black footed nigripes including one from
Sangihe, north of Celebes. However, a few from Celebes appear to have had
the feet blotched black and yellow, much as described by Parkes for Philip-
pine birds. I also note that Siebers (1930: 211) in recording two examples
collected by Toxopeus in Buru in May and June 1921 gives the recorded soft
parts of one as "feet black" and of the other as "feet olive . . . (illegible)."
Thus one of these birds from Buru probably had spotted feet like the Philip-
pine birds. It is thus possible that birds with spotted feet may be a phenotype
which occurs among black footed nigripes rather than a separate geographical
form. New data on Philippine birds is required to throw light upon the fact
that a series from Formosa is black footed like most birds from Wallacea
whilst a few from Wallacea have spotted feet like those so far examined or
described from the Philippines. On the possibility of some individual
variation in foot colour, it should be noted that Hue & Etchecopar (1970: 64)
mention birds with black feet on the south side of the Caspian Sea in the
range of nominate gar^etta.
There is evidence that occasional yellow footed migrant gar^etta reach
Wallacea. One from Menado, north Celebes, and just south of the Philip-
pines, in the Leiden Museum, collected by F. von Faber and unfortunately
not dated, seems to be a migrant nominate gangtta. Another possible example
is one from Ambon collected by Teysmann in 1877 with apparently yellow
feet, though as it had been mounted there remains the possibility of post-
mortem changes.
There remains the question of how far E. gar^etta breeds in Wallacea.
There seem to be no records of breeding colonies and Australian breeding
nigripes are known from ringing recoveries to be strongly migratory, reaching
New Guinea, New Britain and New Zealand. Possibly they also migrate into
Wallacea. Stresemann (1914: 65) reported small flocks as common in Seran
in October and November which could have been such migrants, as could
the two from Buru mentioned above. Such negative evidence is inconclusive
but there seems to be no evidence that E. gar^etta, E. alba or E. intermedia
breeds in Wallacea. All three however are known to breed in heronries in
west Java.
References :
Amadon, D. & Woolfenden, G. 1952. Notes on the Mathews Collection of Australian
birds. The order Ciconiiformes. Amer. Mus. Novit. 1964.
Hue, F. & Etchecopar, R. D. 1970. Les oiseaux du proche et du moyen Orient. Paris:
Boubee.
Parkes, K. C. 1973. Annotated list of the birds of Leyte Island, Philippines. Nemouria 11.
Siebers, H. C. 1930. Fauna Buruana. Aves. Treubia 8 (suppl.): 165-303.
Stresemann, E. 1914. Die Vogel von Seran. Nov. Zool. 21: 25-153.
79
The relationships of Buettikoforella bivittata:
some considerations
by Murray D. Bruce
Received 13th February, 1974
Buettikoferella is the only endemic genus of Timor, with one species, B.
bivittata, considered by Mayr (1944: 171) to be "probably of western origin".
It was originally described in Napothera and subsequently placed in Drymo-
cataphus (=Pellomeum) and Dumetia until separated into a monotypic genus in
1895. The placements did not question its assignment with these Oriental
timaliines. Mayr {pp. cit.\ 158) pointed out that in addition to resembling
these genera, it resembled "many of the grass warblers", such as the Austra-
lasian " Eremiornis, Megalurulus, Cichlornis, and Ortygocichla\ which may be
timaliine and not sylviine, since suggested with allied groups, e.g. Malurus
(Harrison & Parker 1965), but other evidence may not support this (Sibley
1970: 73).
During a visit to Portuguese Timor, in August 1972, I recorded Buet-
tikoferella in open scrub and grass areas of the lowlands, singly or in pairs, on
the ground or perched in low trees and bushes. The flight was short and slow
with dipping, and slight drooping of the long tail, which, on alighting, was
raised and lowered a few times. The behaviour of Megalurulus mariei is ap-
parently similar. It is found singly or in pairs, on the ground, in tall grass and
undergrowth. Its flight is short and direct with dipping, and the tail is thrown
slightly from side to side, cf. Warner (1947: 176), who compared it to
Telmatodytes. Megalurulus shares "a greatly exaggerated tail" with Buettiko-
ferella and Eremiornis carteri. The latter, of similar habits, has been compared
with Stipiturus (McGill 1970: 33).
Cichlornis grosvenori is found on or near the ground in small groups and
clings vertically to bamboo like a wren. Ortygocichla rubiginosa has similar
habits, cf. Gilliard & LeCroy (1967: 205), and cocks its tail upright when on
the ground (R. I. Orenstein pers. comm.). New Zealand's Bowdleri a punctata
may belong here. I observed it singly in swampy grasslands and it has a weak,
undulating flight, drooping its long tail, which it flicks when perched on
grass stems.
Napothera and Pellorneum include primarily terrestrial species of forested
areas, e.g. P. capistratum (Smythies 1968: 420). The skulking Dumetia
hyperythra is found in small flocks in tall grass, undergrowth or on the ground,
and has been compared with Stachyris (Ali & Ripley 1971 : 180).
The strong Asiatic element in Timor's oscines, pointing to close affinities
with timaliine genera for Buettikoferella, may be due to the possible extinction
of the mountain monsoon forest during the Pleistocene, subsequently
"reconstituted only recently" (Mayr 1965 : 41). Similarities of Buettikoferella
with the above 'grass warbler' genera and others, e.g. Graminicola and
Sphenoeacus, may indicate timaliine affinities of some or all of these genera, or
may only reflect convergence through similar feeding adaptations. Additional
observations are needed to determine the extent of possible relationships.
I am grateful to C. W. Benson and C. M. N. White for comments on an
earlier draft of this note.
References :
Ali, S. & Ripley, S. D. 1971. Handbook of the birds of India and Pakistan 6. London: Oxford
University Press.
80
Gilliard, E. T. & LeCroy, M. 1967. Results of the 1958-1959 Gilliard New Britain Expe-
dition 4. Annotated list of birds of the Whiteman Mountains, New Britain. Bull. Amer.
Mus. Nat. Hist. 135: 173-216.
Harrison, C. J. O. & Parker, S. A. 1965. The behavioural affinities of the blue wrens of
the genus Malurus. Emu 65 : 103-1 13.
McGill, A. R. 1970. Australian warblers. Melbourne: Bird Observers' Club.
Mayr, E. 1944. The birds of Timor and Sumba. Bull. Amer. Mus. Nat. Hist. 83: 123-194.
— 1965. The nature of colonizations in birds, pp. 29-47. In H. G. Baker & G. L. Stebbins
(eds.). The genetics of colonizing species. London: Academic Press.
Sibley, C. G. 1970. A comparative study of the egg-white proteins of passerine birds. Bull.
Peabody Mus. Nat. Hist. 32: 1-131.
Smythies, B. E. 1968. The birds of Borneo. 2nd ed. London: Oliver & Boyd.
Warner, D. W. 1947. The ornithology of New Caledonia and the Loyalty Islands. Unpub-
lished Ph.D. thesis. Cornell University, New York, ix -f 232 pp.
Species limits and variation of the New World
Green Herons Butorides virescens and
Striated Herons B. striatus
by Robert B. Payne
Received ) 1 st January, 1974
The Green Herons Butorides virescens and Striated Herons B. striatus have
usually been considered to be distinct species (Hellmayr & Conover 1948,
Eisenmann 1952, Bock 1956, Peters 193 1, Palmer 1962, Wetmore 1965, Slud
1967), though they have been mentioned as possibly conspecific (Hartert
1920, Eisenmann 1955, Parkes 1955). The Green Herons of North America
look different from the Striated Herons of South America (brown-to-
maroon neck in the former, pale grey neck in the latter), but intermediate
birds live in a limited area between these in Panama and in the islands off the
Caribbean coast of South America (Lowe 1907, Van Tyne 1950, Wetmore
1955). The question whether these herons are conspecific has been difficult
to answer because of an earlier shortage of specimens (Hellmayr & Conover
1948: 184) and because wintering northern migrants have not been dis-
tinguished from local birds. A further question about species limits in the
genus has arisen with Harris's (1973) report of possible sympatry with
limited interbreeding of South American B. striatus with the dark Lava
Herons known as B. sundevalli on the Galapagos Islands.
As part of a study of heron systematics (Payne ms.), I examined the 837
New World and about 500 Old World specimens of Butorides available in the
American Museum of Natural History (AMNH), U.S. National Museum of
Natural History (NMNH), Field Museum of Natural History (FMNH), and
University of Michigan Museum of Zoology (UMMZ). I could not differen-
tiate between first-year and older specimens. Birds in juvenile plumage that
had not completed most or all of their postjuvenile moult were excluded from
the colour analysis reported below. Wing lengths (unflattened chord), bill
lengths (unfeathered base of culmen to the tip), and tarsal lengths were
recorded to the nearest mm. Colour of plumage on the cheek and neck was
compared with NMNH specimens selected to represent a smoothly graded
series of nine colours from grey through brown and maroon (Table 1); these
specimens were compared directly with specimens in other museums.
Throughout eastern North America, Green Herons occur with no obvious
geographic differences in colour or size (Parkes 1955, my observations).
81
TABLE i
Colour reference specimens of Butorides striatus
iV<?^ ro/o.w
NMNH cat. no.
Locality
I
grey
263848
Venezuela: Culata
2
grey, tinge brown
423096
Panama: Charco del Toro
3
grey, wash brown
444948
Panama: Rio Indio
4
brownish grey
4001 1 3
Panama: Pese, Herrera
5
greyish brown
448634
Panama: La Jagua
6
greyish red-brown
368472
Colombia: Guajira
7
reddish brown
206343
Panama: Rio Indio
8
purplish brown
316840
Cocos Island
9
dark purplish brown
468699
Panama: Almirante
Green Herons in southwestern North America are larger and include more
birds with paler brown necks than in the east (Fig. I). Overlap in size is little;
informed ringers could distinguish nearly all individuals in areas where both
might winter together, and on this criterion I recognize the western form as
a subspecies {anthony'i) distinct from the eastern birds (virescens). Birds of Baja
California south of 27 latitude are all very dark-necked, more purplish than
brown, and are a distinctive subspecies (fra^ari). In Mexico, little colour
variation occurs but wing length varies from 167 to 190 within Tamaulipas
(considering only "summering" birds, taken between 1 May and 30 Septem-
ber, thereby hopefully excluding any "wintering" birds from more northern
populations). Birds of Central America from Guatemala through El Salvador
and Costa Rica average smaller than the North American sample from Michi-
gan but considerable overlap occurs in wing length and no consistent colour
differences appear. Birds of the West Indies average smaller than the eastern
North American virescens, but, even excluding all winter birds, some overlap
occurs with North American birds and considerable overlap with Mexican
birds. Birds of the Bahama Islands are uniformly smaller and paler than the
North American birds, and though similar in size and colour to a few birds of
the Lesser Antilles, nearly all of the Lesser Antilles birds are darker and many
are larger than the Bahamas birds. The form bahamensis thus appears to be a
valid subspecies. The 1 1 various other described subspecies from some of the
islands and from parts of Central America (including 10 names of Ober-
holser 191 2) are not distinctive.
In Panama, dark-necked virescens occur both on the Caribbean side in the
west (Bocas del Toro) and on the Pearl Islands on the Pacific side farther
east; similar dark birds have been taken on Cocos Island. The Pearl Islands
birds have been described as a subspecies, but similar birds occur on Swan
Island in the western Caribbean, on Puerto Rico, and in Mexico (Tamau-
lipas), all of these in winter. Similar birds are also known from Cuba, the Isle
of Pines, Quintana Roo, and Florida (Bangs 191 5). It seems unlikely that all
of these represent birds that dispersed from the Panama Bocas del Toro or
Pearl Islands population, as the localities are so widespread; probably dark-
necked herons breed as locally differentiated populations or as dark indi-
viduals through several parts of the range of the virescens group. For details of
variation, see Fig. II.
East of 8i° on the Isthmus of Panama both dark-necked and light-necked
herons occur, but all of the dark necked herons (colours 7-9) are wintering
birds and thus all may be migrants from further north. Nests on Barro
Colorado attributed to B. virescens by earlier workers (in Eisenmann 1952: 12)
82
SOUTHWEST
SO BAJA
MICHIGAN
MEXICO
XX X
XXX9 9 9 7 X
9 8 XXX9X X9 X 9 X9X
8 8 8 8 7
L ' \ ®® 7»8 8 ©
7 8 8®®8®8®®7 8(
8 8 8
Qji ©7(7) 7(
© © 8® 7 5® HI
9 5 9 8 7 1 9 8
95859938593 5
58852752169225809 5 8
5533 75928625273995328395887 78
«> ©@ ©5 56 7®6®
5 5 6 6 5(6)5(7)7 7
©888 ©8
©8 8 7@® 8®© 8
! 8 ©8 7 8® ®7® 7
J8 8 8 7® 7@®7 8® 7 8
8©8 7 ®®©7 7®®7®®<3® 8 7®7 8 8 "
SS
® '
1 2® ®2 ® 3 5®
1 2©2
2 12 1
150
170
WING LENGTH, mm
CENTRAL AMERICAN
PANAMA
BAHAMA ISLANDS
GREATER ANTILLES
LESSER ANTILLES
SOUTH CARIBBEAN
TRINIDAD
VENEZUELA
COLOMBIA
190
210
Figure I. Wing length (mm) and neck colour (cf. Table i, except "5" in Fig. 1= "4" or
"5" of Table 1) of Butorides striatus populations in the New World (except most of S.
America). Each number represents one specimen. Circled specimens were taken October to
April and include wintering migrants as well as residents in winter; uncircled figures
represent birds taken from May through September and likely to be breeding birds. Note:
Winter specimens from southwestern U.S., Baja, Panama (see Fig. II), and Colombia
(except wintering northern migrants) are not circled, and wintering Mexican specimens with
wing-length larger than 181 mm are not included. For Southwest, C= Chihuahua; for
Mexico, T=Tamaulipas; for Greater Antilles, PR = Puerto Rico; for Lesser Antilles and
nearby islands, S = Swan Island, V=St. Vincent; for S. Caribbean, V=Espada, Venezuela,
C=Guajira, Colombia, A=Aruba, LR = Los Roques, M= Margarita Id., all others =
Tobago. X indicate a colour darker purple (less brown, more greyish) than colour 9 and is
characteristic of birds of southern Baja.
apparently were identified without collecting the breeding birds ; Van Tyne
(1950) and Wetmore (1965 : 87-88) attribute all breeding birds there to the
intermediate, brownish-grey population. Birds taken from April through
September on the Isthmus west of the Canal Zone (many collected recently
by A. Wetmore) are all intermediate in neck colour between the western
Panama virescens and South American striatus. The sample from Code, 80 km
west of the Canal Zone, averages darker than the nonwinter sample from the
Canal Zone itself (colour scores of 7, 6, 6, 4, 4; vs. scores of 5, 4-5, 4, 3, 3, 3,
2), suggesting a local cline of colour from western Panama-Cocle-Canal
Zone-Colombia, and perhaps some genetic introgression. Birds of the Canal
©,3,®.©,©.®.®,®,®,®,®
2®3,3.®.@.®,®,®,4_5,©®,®-
Figure II. Variation in Butorides herons in Panama. Each number represents the neck
colour of one specimen (cf. Table 1). Circled numbers are winter birds (both residents and
northern migrants), uncircled figures represent probable breeding residents (all May to
September). Note that all non-winter birds in central and eastern Panama (including Canal
Zone) are intermediate in neck score between South American birds (score 1) and western
Panama birds.
Zone have sometimes been regarded as specifically different from B. virescens
and Griscom (1929) described them as a subspecies, B. striatus patens. These
birds were regarded as specifically distinct from B. virescens also by Wetmore
(1965) though he doubted the validity of the subspecies as some South
American birds also have similar brownish-grey cheeks and necks. Hellmayr
& Conover (1948) came to the same conclusion. Van Tyne (1950: 5) con-
sidered these intermediate patens birds to be a race o£B. virescens rather than
B. striatus because the colour of their tarsi were like those of northern
virescens (greenish) rather than southern striatus (reddish) and the colour of
the lores was greenish yellow, not blue as reported for striatus by Griscom
(1929). However, the tarsi of both North American virescens and South
American striatus change seasonally from yellow or greenish to orange or
orange-red in the breeding season (Palmer 1 962 : 41 5, Haverschmidt 1 968 : 1 3),
84
and the lores at least in virescens then change from "dull yellow lime" to "blue-
black" (Palmer 1962: 415). No conspicuous colour differences are obvious in
birds in similar physiological condition. To draw the line between a plumage
colour of birds that here score 4 or 5 (or 5 or 6) and to call this a species
difference seems arbitrary, and in fact the variation in plumage colour is
continuous within these local populations. The Panama birds (excluding
probable northern migrants) share characters with northern and southern
birds and appear to be an intermediate population, probably of hybrid origin
inasmuch as colour is variable within the populations.
Specimens from Cocos Island are few, and most of them are immatures.
Slud (1967) describes both B. virescens and B. striatus from the island, the
latter both the intermediate "patens" birds and birds indistinguishable from
those of South America. Perhaps the birds of extreme plumage colours are
nonbreeding migrants. No birds known to be breeding have been collected
there.
Interbreeding between northern virescens and southern striatus is also
indicated by the wide variation in plumage colour in populations on the
islands off Venezuela and Colombia as well as in coastal populations of these
countries. Birds of Aruba, Curacao, Bonaire, and Los Roques include two
individuals of intermediate colour as are mainland birds from the northern
Gulf of Venezuela (Espada and Guajira). Further east, a bird from Margarita
Island is greyish brown, and the Tobago birds are variable in neck colour
suggesting genetic recombinations of northern and southern herons.
Herons of mainland South America include several specimens intermediate
in colour between northern brown-necked virescens and grey-necked striatus.
Intermediate birds from Venezuela are from Espada, Caicara, Paria, Cumana,
Mt. Duida, and 1800' at San Antonio, all but the last two on the Caribbean
coastal plain. In Colombia, intermediate birds were taken on the Guajira
Peninsula ("laying" bird of colour 6 at Laguna de Tucacas) and in the drain-
age basins of the Cauca and Magdalena Rivers, which flow to the Caribbean.
One bird with colour 4 was taken 500 km upstream from the Cauca mouth, at
Huila, La Plata. Birds of Trinidad, offshore from the mouth of the Orinoco,
are mainly grey-plumaged as are most South American birds. Grey-necked
birds predominate through Colombia except in the Magdalena basin and
Guajira. Also taken in Venezuela and Colombia were birds as large and
brown as the North American virescens •; but all of these were in the northern
winter, and there is no indication of overlap of breeding areas of the dark-
brown-necked and grey birds in South America.
Brownish-grey-necked individuals occur through the rest of South
America as an apparent uncommon colour morph (Table 2). Brownish-
TABLE 2
Variation in neck colour of Butorides striatus of South America*
Colour ol
plumage on
neck:
number of specimens
Country
grey grey
tinge brown
grey
wash brown or
greyish
(1)
(2)
brownish grey (3) (4)
brown (5)
Surinam, Cayenne
2 5
2
1
1
Brazil
128
12
3
Ecuador, Peru
57
9
5
Bolivia
7
3
2
Paraguay, Uruguay
20
3
1
Argentina
21
1
* excluding Venezuela and Colombia (see
Fig. I)
85
necked birds comprise nearly a fifth of the sample in Ecuador and Peru, well
out of the range of the Caribbean drainage region; one with a score of 3 was
taken at Mocori, Chaco, Argentina. The occurrence of these brownish-
necked birds in South America does not necessarily indicate recent genetic
input from the North American populations because the distribution does
not coincide with the southern extent of the range of the wintering northern
migrants. Possibly these forms may reflect an earlier contact of the southern
B. striatus with northern birds, inasmuch as brownish-necked individuals do
not occur in the populations of B. striatus in Africa through eastern Asia.
Griscom (1929: 157) and Amadon (1953: 404) doubted that South American
{striatus) and African (atricapillus) birds could be differentiated. However, in
addition to the absence of brownish-necked individuals, African birds differ
in having a more distinct rufous area on the upper breast, lateral to the lower
neck streaks, and rufous (not grey-brown) lower neck streaks as Hartert
(1920: 156) noted. Other variation among birds within South America
includes colour of the belly; some are whitish, others darker grey, and others
intermediate. The subspecific name "cyanurus" has been applied to southern
birds, which supposedly have paler underparts. I noted considerable varia-
tion within all local populations and much overlap in belly colour in birds
from central Argentina, from northern Argentina and southern Brazil, and
from the more northern regions in South America, and consider it impossible
to determine the locality of individuals on the basis of their underparts. No
significant mean differences in wing length, bill length, and tarsal length
could be determined among these South American populations, they all
overlapped considerably.
The geographic pattern of variation among herons of the Butorides
virescens-striatus complex indicates that they are conspecific. Both in Panama
and on the coasts of Caribbean Colombia and Venezuela and their offshore
islands, a wide range of phenotypes occur intermediate between the maroon-
brown necks of the northern virescens group and the grey necks of the
southern striatus group. This variation suggests secondary contact of two
forms previously isolated geographically that have since interbred. These
populations, lying mainly between 8° and io° N latitude, are probably
sedentary and receive limited gene flow from the northern and southern
populations inasmuch as intermediates outnumbered the extremes in central
and eastern Panama and on the off-shore Caribbean islands near South
America. However, the wide range of neck colour and the presence of some
dark-necked or grey-necked birds suggests some possible limited gene flow
resulting from immigration, perhaps involving occasional local breeding of
northern immigrants that may remain on their wintering grounds with the
local population. The small size of the southern Caribbean birds may reflect
past inter-breeding of northern South American birds with the small birds
of the West Indies (cf. Fig. I). Because the birds of Panama and the southern
Caribbean are variable and combine the differing character states of their
neighbouring populations, it is unnecessary to recognize any subspecies
within this region.
The species name to be used for the combined complex appears to be
Butorides striatus. Linnaeus described both striatus and virescens in the same
edition (10th, 1758) on the same page (p. 144) of "Systema Naturae." Parkes
(1955) suggested that Lowe's 1907 treatment of the form "robinsoni" from
Margarita Island was the first revision to recognize one of these forms as
conspecific with the other. Lowe referred to two specimens as " Butorides
86
virescens robinsoni" on p. 554 but twice on p. 555 named them "B. robinsoni"
and he contrasted this form to the South American birds, which he called
"B. striata." Lowe did not state that striatus was conspecific with virescens,
though such an act would be required {International Code 1961) for purposes
of establishing a priority of names. I do not think that Lowe's attempts to
describe two specimens with reference to no other forms designated more
specifically than "B. striata" and "B. virescens" constitutes a first revision, as
the Code Article 24 (a) (i) makes it clear that "The expression 'first reviser' is
to be rigidly construed." Lowe appeals to have regarded "'robinsoni" itself
as being as much of a species as he did virescens and striatus. On the other
hand, Hartert (1 920 : 1 249-1 2 5 1) described and revised the group on a world-
wide basis; he listed virescens as a subspecies of Butorides striatus (p. 1250). As
Hartert's usage of B. striatus appears to be the first definite decision that the
forms were conspecific, the first reviser principle requires that the species
name to be used is Butorides striatus.
The Lava Heron of the Galapagos Islands has long been considered a
species Butorides sundeva/fi, though Hartert (1920: 1250) regarded it as a sub-
species of B. striatus. "Typical" adult specimens are dark grey, with blue-
green lanceolate back plumes and a suggestion of pale streaks on the throat;
the feet are red as in other breeding herons of the complex. Harris (1973:
267-268) has called attention to the occurrence of B. s. striatus-likc birds on
the islands and to specimens intermediate in appearance between sundevalli
and striatus. The 50 specimens I have examined (NMNH, AMNH, FMNH,
and 8 from the California Academy of Sciences) show all possible intermediate
plumage colours (Table 3). Most of the paler birds differ from the South
TABLE 3
Colour variation in Butorides striatus of the Galapagos Islands
No. birds Plumage colour
13 underparts blackish, unstreaked or with white edges of feathers on
centre of throat.
13 (4) underparts dark grey, nearly blackish, unstreaked except on throat; or
throat grey, unstreaked (N = 2).
11 (1) underparts dark grey, belly lighter grey; sides of neck and cheeks
blackish to dark grey ; throat streaked with white edges.
3 (1) underparts grey, darker than South American B. s. striatus; sides of
neck and cheeks grey, below streaked white and grey-brown to upper
breast (feathers of midline mainly white).
2 (2) underparts light grey, like South American B. s. striatus', neck streaked
white and grey-brown to upper breast; may have some rufous on
upper breast but most birds have more grey-brown (less rufous) than
B. s. striatus.
Numbers in parenthesis are a selected non-random sample of CAS birds. These numbers are
not included in the first column. All of the plumage colour classes appear to intergrade
with the next lighter and darker colours.
American B. s. striatus in having less rusty colour on the upper breast; the
neck streaks of the Galapagos birds are grey-brown and the sides of the upper
breast are grey, with no more than a trace of rufous, but otherwise the pale
birds are inseparable from the South American birds. The difference in
amount of rusty colour on the neck and breast indicates that the pale birds of
the Galapagos are mainly local birds, not visitors from the mainland. Both
the paler and the darker birds are widespread, and both occur together on
87
some islands. The high proportion of intermediate birds indicates consider-
able successful breeding and genetic recombination. Perhaps some differen-
tiation of the herons occurred on different islands with subsequent inter-
island reinvasion and interbreeding, or more likely the islands were invaded
successfully by more than one wave of immigrants from South America in
the past: either history might account for the variation among the Galapagos
herons. The Galapagos herons are variable, but nearly all adults are separable
from adults of the continental populations. The Lava Heron is probably best
considered a distinct subspecies, B. striatus sundevalli, at least until studies on
breeding pairs have been made in the islands.
Acknowledgements : For loaning or providing access to their material I thank
W. E. Lanyon, J. Farrand, and J. J. Morony (American Museum of Natural
History), L. C. Binford (California Academy of Sciences), R. L. Zusi (Nation-
al Museum of Natural History), and M. A. Traylor (Field Museum of Natural
History). I thank the F. M. Chapman Memorial Fund for support. L. C
Binford, E. Eisenmann, K. C. Parkes, R. W. Storer, and A. Wetmore read
the manuscript and offered helpful criticism. The figures were drawn by
Patricia J. Wynne.
References :
Amadon, D. 1953. Avian systematics and evolution in the Gulf of Guinea. Bull. Amer. Mus.
Nat. Hist. 100: 393-452.
Bangs, O. 191 5. Notes on dichromatic herons and hawks. Auk 32: 481-484.
Bock, W. J. 1956. A generic review of the family Ardeidae (Aves). Amer. Mus. Novit. 1779.
Eisenmann, E. 1952. Annotated list of birds of Barro Colorado Island, Panama Canal Zone.
Smiths. Misc. Coll. 117(5): 1-62.
— 1955. The species of Middle American birds. Trans. Linn. Soc. N.Y. 7.
Griscom, L. 1929. A collection of birds from Cana, Darien. Bull. Mus. Comp. Zool. 69: 147-
1190.
Harr s, M. P. 1973. The Galapagos avifauna. Condor 75 : 265-278.
Hartert, E. 1920. Gattung Butorides Blyth. In Die Vogel der palaarktischen Fauna, 1912-1921,
vol. 2: 1249-125 1. Berlin: Friedlander.
Haverschmidt, F. 1968. Birds of Surinam. Edinburgh: Oliver & Boyd.
Hellmayr, C. E. & Conover, B. 1948. Catalogue of birds of the Americas and the adjacent
islands. Field. Mus. Nat. Hist., Zool. Ser. 13, 1(2).
Lowe, P. R. 1907. On the birds of Margarita Island, Venezuela. Ibis (9)1 : 547-570.
Oberholser, H. C. 1912. A revision of the subspecies of the Green Heron [Butorides virescens
(Linnaeus)]. Proc. U.S. Nat. Mus. 42: 529-577.
Palmer, R. S. (ed.). 1962. Handbook of North American birds, vol. 1. New Haven, Conn.:
Yale Univ. Press.
Parkes, K. C. 1955. Systematic notes on North American birds. 1. The herons and ibises
(Ciconiiformes). Ann. Carnegie Mus. 33: 287-291.
Payne, R. B. In ms. Family Ardeidae. In Peters Check-list of birds oj the world, vol. 1, ed. 2.
Ed. E. Mayr. Cambridge, Mass. : Harvard Univ. Press.
Peters, J. L. 193 1. Check-list of birds of the world, vol. 1. Cambridge, Mass.: Harvard Univ.
Press.
Slud, P. 1967. The birds of Cocos Island (Costa Rica). Bull. Amer. Mus. Nat. Hist. 134(4):
261-296.
Van Tyne, J. 1950. Bird notes from Barro Colorado Island, Canal Zone. Occ. Pap. Mus.
Zool. Univ. Mich. 525.
Wetmore, A. 1965. The birds of the Republic of Panama, part 1. Smiths. Misc. Coll. 150.
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Published by the BRITISH ORNITHOLOGISTS' CLUB and printed by
The Caxton and Holmesdale Press, 104 London Road, Sevenoaks, Kent.
Bulletin of the
British Ornithologists' Club
t
2 6 SEP 1974
Edited by
HUGH F. I. ELLIOTT
Volume 94 No. 3
September 1 974
Special General Meeting
Notice is hereby given that a Special General Meeting of the Club will be
held at the Cafe Royal, 68 Regent Street, London, W.i. on Tuesday, 19th
November 1974 at 6 p.m. to consider and if deemed fit to pass the following
resolution which will be proposed by the Committee as a special resolution
in the terms of Rule (14) of the Club's rules: —
"That with effect from 1st January 1975 the annual subscription for members
shall be £3 • 50 and that Rule (4) shall be amended accordingly".
By order of the Committee
RONALD E. F. PEAL
Honorary Secretary
Note: Since the annual subscription for members was raised to £2 10s. with
effect from 1st January 1971 printing costs have risen over 48%. Most of
the Club's expenditure is on printing and, with continuing inflation, an
increase in the subscription rate is now essential to avoid the likelihood of a
heavy financial deficit in the forthcoming year.
Bulletin of the
BRITISH ORNITHOLOGISTS' CLUB
Vol. 94 No. 3 Published: 20 September, 1974
The six hundred and eighty-eighth meeting of the Club was held at the Cafe
Royal, 68 Regent Street, London, W.i., on Tuesday, 14th May 1974 at 7 p.m.
Chairman: Mr. Peter Hogg; Present 18 members and eight guests.
The Speaker was Mr. D. T. Holyoak, who addressed the Club on Orni-
thological exploration and conservation in the Cook islands. He exhibited
the skins of seven new forms from the southern Cook Islands, descriptions
of which will be published shortly.
The six hundred and eighty-ninth meeting of the Club was held at Imperial
College, Exhibition Road, London, S.W.7. on Tuesday, 2nd July 1974 at
6.45 p.m.
Chairman: Mr. Peter Hogg; Present 16 members and 11 guests.
Mr. Richard Blindell spoke on his current investigations into waders and
wildfowl on the Essex coast and in the Thames estuary. After supper he was
followed by Mr. Ian R. Deans, who addressed the Club on the results to date
of work he is undertaking on gull ecology in the Thames estuary.
Duetting in Hypergerus atriceps and its Taxonomic
relationship to Etninia lepida
by L. G. Grimes
INTRODUCTION
Hypergerus Reichenbach, 1850, Syst. Av., pi. 54 (9), with type by monotypy
Moho atriceps Lesson, 183 1, Traite Orn., p. 646, is a monotypic genus in the
Passeriforms of uncertain affinities and so is Eminia Hartlaub, 1881, Proc.
Zool. Soc. Lond, p. 625, with type by monotypy Eminia lepida Hartlaub with
the same citation. Recent authorities (White i960, Hall & Moreau 1970)
hesitantly follow Chapin (1953) in assigning both genera to the warblers
Sylviidae. Both species are found in similar habitats, Hypergerus in West
Africa and Eminia in East Africa (White i960), and Chapin found so many
similarities between the two in the field that he surmised that the nest of
Hypergerus when found would probably be very similar to that of Eminia
(Chapin 1953, Van Someren 1956). This has been confirmed by Lang (1969).
Lanyon (1969) has emphasised that vocal characteristics may reveal close
relationships between avian species which may not be discernible on the
basis of morphology alone, and also may enable the recognition of groups of
allopatric forms that comprise a superspecies. This approach has not been
applied to species that are both morphologically distinct and allopatric, and
placed in separate genera. Hypergerus atriceps and Eminia lepida (Hall &
Moreau 1970) fit this category and if they are more closely related than their
placement in separate genera suggests, then, following Lanyon's work, one
might expect to find some similarities in their song structures.
The present study was stimulated through the review by Thorpe (1972) of
duetting in birds. This is well known in Eminia (Chapin 1953, Van Someren
1956). Although Hypergerus duets regularly this has only recently been
recognised. The occurrence of duetting in both species suggested that
similarities, if any, in their duet structure might provide evidence for a close
relationship between them. In this paper the vocal characteristics of the two
species and their duets are compared and described, that for Hypergerus for
the first time. Other comparative data on their biology is reviewed to support
the thesis of this paper that the two species are congeneric.
MATERIALS AND METHODS
Two recordings of Hypergerus, involving the same birds, were made on
28 August 1972 in an area of dry thicket on an inselberg on the Accra Plains
in southern Ghana. The time length of the recording was four minutes and
includes 40 duet sequences. A Philips recorder (Model 2204) was used with
the microphone placed at the focus of a parabola (o-6 m diameter). This
recording was dubbed at 3-75 in/sec using a Philips recorder (Model 4307)
on to a master tape which was used for all subsequent analysis with the Kaye
sonograph in the wide band mode.
Two recordings of Eminia made by the late M. E. W. North in Kenya
were obtained from the library of sounds of the Laboratory of Ornithology
of Cornell University. The first was recorded on 17 and 18 December 1953
at Gituru (i° 10' S, 3 6° 50' E), and although it is just under seven minutes
in length only nine duet sequences occur : part of this recording is found on
the record "Voices of African Birds" (North 1958). The second recording
of a single male was obtained on 6 September 1963, at Lake Nakuru (o° 15' S,
3 6° 05' E), and lasts for just over one minute.
RESULTS
In both species one bird, presumably the male, dominated the singing and
always initiated the duet sequences. Although there are marked differences
5
H2 4 -
5 -
Hz 4 A
B
Fig. 1
v l
- c
**
r^s.
j, s ^
A
L
\
9O
Table i. The analysis into phrases of the song of two duet sequences of Hypergerus
atriceps recorded in southern Ghana. The bar over the syllables indicates the occurrence of
duetting and the underlined sections have been reproduced as spectograms.
K D D BAC8AAACBAACBAA
DUET
SEQUENCE (I)
E A C B C D D
CBAACBCBCBAC OBACBCBCBCDBACDBCB
I 6 Seel
CDBCDBACBAACDL
I 2 2Se
CBACBCBCODBACD
F B A A C
I 4«S«c
I 1 I I I I I 1 I I 1 I I I
in the songs of the males, the second component which forms the duet is
extremely similar in both species and is presumed to be given by the female.
The song of the male Hypergerus consisted of mixed combinations of song
phrases (Figure i and Table i) several of which are either slight modifications
or incomplete forms of others: compare phrases B and N with A, phrases
D, E and L with C, and I with H. Some phrases occur more frequently than
others (A, B, C, H, G, and I), and in some sections the song sequence is a
repetition of one phrase (G, H and I). The major drawback of the presen-
tation in Table i (and Table 4) is that the time interval between the ending
Table 2. Typical values of the ratio of the time difference between the ending of a song
syllable and the beginning of the next to the time difference between notes within the
syllables of the song of Hypergerus atriceps. The time difference is in mm where 132 mm =
1 sec.
Time difference ratio
16
16
20
19
16
16
19
22
12
14
13
Syllable
A
B
C
D
E
F
G
H
I
J
K
12
12
15
15
14
18
13
21
1 2
5
Fig. 2
UIHI
^MMMMMMflflO ^ p>
5 -
Hz 4
1 ;
MMAMflMMMAIIfl^Q y
TIME
91
Table 3. The maximum trill rates (notes/sec) occurring in four examples of the female
component of the duets of Hyper gerus atriceps and Eminia lepida.
MERGER US
EMINIA
11 -7
17 -o
12-4
18 -2
10 -4
18-3
11 7
17-8
N\\\\\\
Fig. 3
\\\\\\\\\\\\\
/ ••••••••••/'••
uunnu
— "inminmtif
G - H
WWW
» * * *%%>*sm*
\:\\\\
\\\
1 //////
^NWSWW
M
A A
92
of a phrase and the beginning of the next cannot be represented. For
Hypergerus these time intervals are only slightly greater than the time intervals
between the notes in a phrase (Table 2). The result is that to the human ear
the male song sounds regular and flowing.
The female component that forms the duet is a trill (Figure 2, Table 3)
each note of which has a similar frequency/time profile. Generally, there
is a slight increase in the trilling rate at the beginning of a duet and it then
either remains constant during the rest of the trill or decreases as the trill
component ends (Figure 2).
Only one recorded sequence of Eminia contained a combination of
different song phrases. The rest were made up of repetitions of the same phrase
(Figure 3 and Table 4). The time intervals between the song phrases vary
from one to four seconds or more and are very much greater (approximately
20 x) than the time intervals between the notes making up each song
phrase. They are also longer than the lengths of the song phrases. As a result
the male song is of a much different tempo and rhythm than the song of the
male Hypergerus. However, the duet component is a trill (Figure 4), and in
structure and tempo it is remarkably similar to that of Hypergerus (Table 3).
Table 4. The analysis into phrases of the song of two male Eminia lepida recorded in
Kenya by the late Mr. Myles-North. The recordings are not continuous and the breaks are
indicated by the vertical bars. The bar over the syllables indicate the occurrence of duetting
and the underlined sections have been reproduced as spectograms. Two sections indicated
( ) have not yet been analysed because of background calls from other species.
Recording at aA^aIaI | i3 Hfl|jj|cCCc|c i £CCc|lJr>DDDi>|DD ] £DD|EEKEEE|
Gituru. I . 1 I I 1 _ _ 1
EEBUES ^Pf??f?t' GGGG.GGGG HH|H HHHHH H 1IHH II I I
Section from
record "Voice I
of African I _I I iJC J J J K LM 1: bj b U
Birds" '
Recording at .
Lake Nakuru N N N N N N NNNNNNPPPPPPP
Fig- 4
uu urn
TIME
OTHER COMPARATIVE DATA
In both species the sexes are alike and the juveniles closely resemble the
adults, more so in Eminia than Hypergerus. Following Chapin's key for the
genera in the Sylviidae (Chapin 1953: 241) both birds have similar values
93
(°'33 — o-66) for the ratio of the lengths of the two outermost primaries,
they have a similarly shaped bill without a hooked tip, and both have rela-
tively long tails. In both species males are larger than females. Details for
Hypergerus are given in Table 5. Chapin (1953: 308) gives no details for
Table 5. The physical dimensions (in mm) of Hypergerus and Eminia. The number of
specimens used are given in brackets after the sex symbol.
Hypergerus
Source
Sex
Wing
Tail
Tarsus
Bates (1930)
6* (4)
84-87
94-102
27-30
$(3)
77-82
83-94
Bannerman (1936)
6* (5)
83-88
91-99
28-30
9(8)
76-82
Eminia
81-92
27-28
Reichenow (1905)
?
65-71
5 5-65
23-25
Eminia but mentions that males are consistently larger than females. The
measurements for Eminia by Reichenow (1905 : 615) are included in Table 5
for comparison, but unfortunately he neither includes sample size nor dis-
tinguishes between the sexes.
Both species breed in the wet season and both are probably double
brooded. This is certain for Eminia (Van Someren 1956: 339) but not yet
certain for Hypergerus ', although the observations of Lang (1969) suggest this.
Old nests may be reconditioned and used again in both species.
The clutch size is usually two eggs for Eminia, sometimes three. The two
clutches known for Hypergerus had two and three eggs respectively. The eggs
of Eminia are either pure white or pale blue or else have in addition some
dark purple brown (reddish brown) spots. The two eggs of Hypergerus
found by Lang (1969) were pale blue with reddish brown spots and blotches
fairly densely distributed at the blunt end and scattered lightly over the rest
of the egg. In both species the shape of the eggs is elongated oval ; the ratio
of length to breadth for Eminia being in the range of 1.44 to 1.59 and for
Hypergerus 1.48 (only one egg measured).
The nests of the two species are remarkably alike, both resembling a large
untidy sunbird nest and are not unlike those of the eurylaemid Smithornis
capensis (R. K. Brooke in iitt.). The nest site and materials used in construction
are also similar. The details that follow are taken from Van Someren (1956:
377) for Eminia and from Lang (1 969 : 1 27) for Hypergerus. In Eminia the nest
is an oversized untidy sunbird nest and is suspended from the end of a thin
branch, usually overhanging water or a stream bed or else a small open space.
The nest has a slight porch and the entrance is on the side near the top.
Strips of bark fibre, grass and tendrils and long strands of moss make up the
outside structure of the nest but all are loosely interlocked and the whole is so
untidy that it looks like a mass of debris caught up on a thorny spray, or a
collection of flotsam brought down in a flood and caught up by a over-
hanging branch. Finer fibres are used as a lining and a few feathers may be
included.
In Hypergerus both nests were suspended from a hanging tendril overhang-
ing a stream. The side entrance was situated near the top of the nest and there
was a slight overhanging porch. Below the chamber of the nest hang an
untidy mass of ragged grass ends. One of the nests was made entirely of
fairly broad strips of dry grass which had collected as flotsam. The second
94
nest had, in addition, a few pieces of dried plant. Both nests had a lining of
much finer grass.
The other genera in the Sylviidae have in general quite different nests and
there appear to be only two species which have nests bearing any resemblance
to a sunbird's. The first is Sylvietta virens (Chapin 1953: 257, 258) and the
second is Apalis pulchra (Chapin 1953: 303). In both cases, however, the
materials used are different from those used by Eminia and Hypergerus.
DISCUSSION AND CONCLUSION
Hall and Moreau (1970) describe the basic distributional patterns that may
have given rise to past speciation in African passerines. They further des-
cribe what they call secondary distributional patterns in order to consider
the effect of subsequent climatic changes on these species. Their secondary
pattern relevant to the present discussion is one where the isolated species
remain so and continue to diverge, even to a level at which they might well
be classified as separate genera. Such a taxonomic situation could under-
standably arise when two such isolated species, one say in West Africa and
the other in East Africa, are collected and described by different people or by
a person such as Hartlaub who although he described Eminia was more
familiar with West African birds than East African birds. This is especially
the case when comparative data are lacking. Hall and Moreau (1970) list five
examples in which they believe this type of isolation and divergence have
occurred. These involve the Sunlark and allies (Galerida modesta), the
Moustached Warbler and allies (Sphenoaecus men/a/is), the Black-headed
Stream Warbler and allies [Bathmocercus cerviniventris), the Barred Warbler
and allies {Cameroptera fasciolatd), and the Olive Sunbird and ally {Nectarinia
olivaced). Such a situation also seems to apply to r/ypergerus and Eminia , first
collected and described respectively in 1831 and 1881. It appears that J. P.
Chapin is the only person who has had the opportunity of comparing them
both in the field. The impression he gained at Lagos, Nigeria, was that
Hypergerus resembled an overgrown warbler. Although the beak, wings and
tail were much more elongated in Hypergerus than in Eminia Chapin (1953 :
309) considered there were many points of similarity but does not list them.
The similarities between the breeding behaviour of Hypergerus and Eminia,
especially that of the nest site and structure, are much greater than would be
expected if they are correctly placed in separate genera. Such similarities are
rather to be expected between species placed in the same genus. On this
evidence alone, therefore, there is some justification for recommending that
they both be placed in one genus. This would appear even more justified now
that it is known that both species duet and that the form of the "female"
component in the duet is the same in both species. This approach does as-
sume that the vocal characteristics being compared remain unchanged.
Lanyon (1969) found such evidence for this in the dawn songs of the tyrannid
flycatchers he studied but this is not available in the present study as only two
species are involved. Nevertheless, it is proposed that Hypergerus a triceps and
Eminia lepida be considered congeneric and be placed in the genus Hypergerus,
the senior of the two names. It is also recommended that the vernacular name
should be the Oriole Warbler adopted by Mackworth Praed & Grant (1973 :
312).
SUMMARY
Hypergerus atriceps and Eminia lepida are placed in monotypic genera in the
warbler family Sylviidae. They are allopatric species, Hypergerus being found
95
in West Africa and Eminia in East Africa. The nest of Hypergerus has recently-
been described and resembles in every way the nest of Eminia. Their duet
structures are described and compared. Although the "male" contributions
are different the "female" components of the duet are very much alike.
These similarities, taken together with their present-day distributions, sug-
gest that they had a common original population whose distribution was
split. The isolates then diverged to produce two species so morphologically
distinct as to suggest that they should be placed in different genera and even
different families.
On the basis of the similarities in their nest, duet structures and some
morphological proportions it is proposed that they be placed in one genus
Hypergerus.
ACK NO WLE DGEMENTS
I thank Dr. M. Lock, L. Lock and R. K. Brooke for their helpful comments
at various stages of the writing of this paper. Also I thank D. W. Lamm for
making it possible for me to obtain the recording of Eminia from Cornell
University.
References :
Bannerman, D. A. 1936. The birds of tropical West Africa. Vol. 4. London: Crown Agents.
Bates, G. L. 1930. Handbook of the birds of West Africa. London: Bale & Danielsson Ltd.
Chapin, J. P. 1953. The birds of the Belgian Congo. Vol. 3. Bull. Am. Mus. Nat. Hist. 7 5 A.
Hall, B. P. & Moreau, R. E. 1970. An atlas of speciation in African passerine birds: London.
Lang, J. R. 1969. Nest and eggs of the Moho or Oriole Babbler. Bull. Nig. Orn. Soc. 6:
127-128.
Lanyon, W. E. 1967. Revision and probable evolution of the Myiarchus flycatchers of the
West Indies. Bull. Am. Mus. Nat. Hist. 136: 392-370.
— 1969. Vocal characters and avian systematics. In Bird Vocalisations (Edit. R. A. Hinde).
Cambridge University Press.
Mackworth-Praed, C. W. & Grant, C. H. B. 1973. Birds of West Central and Western Africa.
Vol. 2. London: Longmans.
North, M. E. W. 1958. Voices of African Birds (Record). Ithaca, New York.
Reichenow, A. 1905. Die Vogel Afrikas. J. Neumann, Neudamm.
Thorpe, W. H. 1972. Duetting and antiphonal song in birds: its extent and significance.
Behaviour Suppl. XVIII Leiden : E. J. Brill.
Van Someren, V. G. L. 1956. Days with birds: studies of habits of some East African species.
Chicago: Natural History Museum Press.
White, C. M. N. i960. A check list of the Ethiopian Muscicapidae (Sylviinae). Part 3.
Occasional papers of the National Museums of Southern Rhodesia. 26B: 695-738.
Status of the Great Shearwater in Trinidad,
West Indies
by Charles T. Collins <& Elisha S. Tikasingh
Received 1 yth fanuary, ip/4
The Great Shearwater Puffinus gravis was first recorded for the island of
Trinidad by ffrench & ffrench (1965) on the basis of the skull of a bird found
dead on an east coast beach in July i960. Since this skull could not be found
at the time of a review of the shearwater records for Trinidad and Tobago
(Collins 1969) it was felt that some caution should be exercised in including
the Great Shearwater in the avifauna of Trinidad until more conclusive
specimen material had been obtained. The preservation of five specimens of
this species obtained during June 1971 and 1972 now fully confirms the
occurrence of the Great Shearwater in Trinidad waters. The details are as
follows.
96
On 10 June 1971, an employee of the Trinidad Regional Virus Laboratory
noticed a large number of dead and dying sea birds on Manzanilla Beach
on the east coast of Trinidad. Unfortunately these were not reported until 18
June at which time the junior author went to the scene and obtained two
birds. One of them was already dead and infested with maggots, while the
second, although badly weakened, was still alive. These birds were later
identified by the senior author as Great Shearwaters and subsequently
prepared as museum specimens, now housed in the collections of the
Trinidad Regional Virus Laboratory (TRVL 15 891) and the American
Museum of Natural History, New York (AMNH 804303; formerly TRVL
15892). At the time these specimens were obtained on 18 June, a rough
estimate of forty additional dead and decomposing shearwaters, presumably
all Great Shearwaters, were observed by the junior author on Manzanilla
Beach.
During June 1972, three additional specimens of Great Shearwaters were
obtained from Trinidad beaches. Single dead or dying birds were picked up
on Matura Beach on 1 June by P. R. Bacon and on 10 June by G. Gibbs. The
third bird, found on Matura Beach by Bacon on 12 June, was alive but unable
to fly. These birds were later skinned by Bacon and R. rTrench and sent to
the senior author for final preparation. Of the three, one specimen (CTC 974)
has been presented to the collections of the British Museum (Natural History)
(1 974.2.1) and the remaining two (CTC 975 and 976) have been added to the
collections of the Trinidad Regional Virus Laboratory and the American
Museum of Natural History (AMNH 810475).
Dead and dying Great Shearwaters were also noted on Matura Beach on
10, 15 and 25 June 1973. Two specimens are preserved in the collections of
the University of the West Indies, St. Agustine, Trinidad (P. R. Bacon in
litt.).
The five specimens from 1971 and 1972 (four males, one female) were in
fresh, nearly unworn, plumage showing no signs of moult of body or flight
feathers. All showed a reduced amount of white on the rump and a larger
brownish patch on the belly as compared with the large AMNH series. The
reduced rump patch is due to the upper tail coverts being strongly mottled
with brownish grey and having the white confined to the terminal edges.
Adult Great Shearwaters leave the breeding colonies in the South Atlantic
during early April (Rowan 1952) and "winter" in the North Atlantic during
which time (July-August) the annual moult of flight feathers begins (Palmer
1962). On the annual northward flight post-breeding adults would be in a
recognisably worn plumage. Even on the breeding grounds the adults are so
worn as to appear paler than the fresh plumaged young (Rowan 1952: no).
Thus it seems reasonable to assume that the fresh plumaged birds recovered
in Trinidad in June are all young of the year which had only recently left the
breeding grounds. No mention has been made by previous authors of the
reduced white on the rump of juveniles although it has been suggested
(Bannerman 1959) that the belly patch is more prominent in juveniles than in
adults. Further observations made on the breeding grounds are needed to
confirm these plumage differences.
As noted earlier (Collins 1969) the appearance of the Great Shearwater in
Trinidad waters is not unexpected in that there seems to be a regular north-
ward movement of this species in the western Atlantic and eastern Caribbean
(Voous & Wattel 1963; Metcalf 1966; Rowan 1952; Hawkins 1963). More
recently (19 June 1971) a specimen of this shearwater was obtained further
97
to the west in the vicinity of Isla Carenero, Islas Los Roques, Venezuela
(Phelps 1972). Since only juveniles have been obtained in Trinidad to date it
suggests a separate migratory route for adults and juveniles as has been
previously noted for the Manx Shearwater Puffinus puffinus (Post 1967).
However, further observations are needed to exclude the possibility that the
adults which leave the breeding grounds (early April) one or two months
before the young (mid-May to early June) (Rowan 1952) pass through this
area at an earlier date.
Mallophagan ectoparasites were abundant on all specimens and ticks were
noted in the ear passages of several. The lice from the two 1971 specimens
were identified by Theresa Clay as Halipeurus gravis Timmermann 1961,
Trabeculus hexacon sens. lat. (Waterston 1914) both of the family Philopteridae
and Austromenopon paululun (Kellog & Chapman 1899) of the family Meno-
ponidae. A single female specimen of a species of Saemundssonia was also
collected from one bird. The ectoparasites from the later specimens have not
yet been identified.
Mass mortalities, particularly of juveniles, similar to that noted on Man-
zanilla Beach in June 1971, have been previously recorded for this and other
species of shearwaters. The most recent account is that of Watson (1970)
documenting the death of hundreds of mainly juvenile Great Shearwaters
along the east coast of the United States in early June 1969. The causes of
juvenile mortality are unknown although storms at sea, bacterial or viral
infections, and the effects of accumulations of organochlorines have all been
suggested. None of these effects could be implicated as the cause of the deaths
recorded by Watson (1970). Similarly, only low levels of polychlorinated
biphenyls were recorded in two of the Great Shearwaters recovered in
Trinidad in 1972 (Bourne & Bogan 1972) and no arbovirus was isolated from
a single bird tested in 1971. The ticks recovered from these birds were not
tested for arboviruses. It should be noted that two of the males obtained in
1972 weighed 385 and 447 g or 44 to 51 percent of the average weight of
adult males (Hagen 1952, in Palmer 1962). The birds reported by Watson
also thin, devoid of fat, and were only about 60 percent of normal adult
breeding weight. Such low weights are probably associated with the cause of
death rather than age as young shearwaters outweigh adults as nestlings. The
actual cause of death however remains a mystery.
The senior author's field work in Trinidad was supported by grants from
the Frank M. Chapman Memorial Fund of the American Museum of Natural
History. Manuscript preparation was aided by a summer faculty fellowship
from the California State University, Long Beach, Foundation. We are
grateful to P. Bacon, R. ffrench and G. Gibbs for contributing the 1972
specimens and their field notes to this study, and to Theresa Clay for identi-
fying the Mallophaga.
References :
Bannerman, D. A. 1959. The birds of the British Isles 8. London: Oliver & Boyd.
Bourne, W. R. P. & Bogan, J. A. 1972. Polychlorinated biphenyls in North Atlantic
seabirds. Marine Pollution Bull. 3: 171-175.
Collins, C. T. 1969. A review of the shearwater records for Trinidad and Tobago, West
Indies. Ibis 111: 251-253.
ffrench R. P. & ffrench, M. 1966. Recent records of birds in Trinidad and Tobago. Wilson
Bull. 78: 5-1 1.
Hawkins, R. S. 1964. Northward passage of Great Shearwaters in the North Atlantic in
May 1963. Sea Swallow 16: 64-65.
Metcalf, W. G. 1966. Observations of migrating Great Shearwaters Puffinus gravis off the
Brazilian coast. Ibis 108: 138-140.
98
Murphy, R. C. 1936. Oceanic Birds of South America. New York: Amer. Mus. Nat. Hist.
Palmer, R. S. (ed.) 1962. Handbook of North American Birds 1. New Haven, Conn.: Yale
University Press.
Phelps, W. H. Jr. 1972. Adiciones a las listas de aves de Sur America, Brasil y Venezuela y
notas sobre aves Venezolanas. Bo/. Soc. Vene^. Cienc. Nat. 30: 23-40.
Post, P. R. 1967. Manx, Audubon's and Little Shearwaters in the northwestern North
Atlantic. Bird-Banding 38: 278-305.
Rowan, M. K. 1952. The Greater Shearwater Pnffinus gravis at its breeding grounds. Ibis
94: 97-121.
Voous, K. H. & Wattel, J. 1963. Distribution and migration of the Greater Shearwater.
Ardea 51 : 143-157.
Watson, G. E. 1970. A shearwater mortality on the Atlantic coast. Atlantic Nat. 25 : 75-80.
Butastur and Buteo east of Wallace's Line
by C. M. N. White
Received jist May, 1974
Brown & Amadon (1968, Eagles, hi air ks and Falcons of the World: 540) give
the range of the Rufous-winged Buzzard Eagle Butastur lirenter as extending
east of Wallace's Line in Indonesia to Celebes, Banggai and Sula Islands. This
requires amendment. All the evidence indicates that this species is or was
common in the south west peninsula of Celebes north to its base at Oeroe
and Palopo (there are eight skins in the Zoological Museum, Bogor, the most
recent dated 7 November 191 2, according to Lord Medway in litt.), but it is
not recorded elsewhere in the island. On failing to trace any published
records from Banggai and Sula, I asked Dr. Amadon why these islands were
included in the range. He informed me that the source was a manuscript
prepared by the late Dr. E. Stresemann in anticipation of a new edition of the
first volume of Peters' Check List of Birds of the World y and that he did not
know the basis of Stresemann's statement. Stresemann set out the range of
B. Uventer in 1940 {foam. Ornith.: 477) but does not include these localities.
Nor does van Bemmel include the species in his list of birds of the Moluccas
(1948, Treubia, 19: 323-402) which covers Sula but not Banggai. It seems
certain that the locality Banggai must be an error, for Dr. S. Eck of the
Staatliches Museum fur Tierkunde, Dresden, who is shortly to publish a
list of the birds of Peleng and Banggai, has kindly informed me that he knows
of no record of B. Uventer from these islands. There are no specimens from
Sula in the American Museum of Natural History, New York (Mrs. M.
LeCroy in lift.) nor in the Leiden Museum (Dr. G. F. Mees in litt.). J. J.
Menden collected at Taliaboe in 1938, but his collection, now at the Museum
of Comparative Zoology, Harvard, does not include this species (Dr. R. A.
Paynter in litt.). Unless this note elicits new information Sula should therefore
also be deleted from the range.
Mentioned by Stresemann (1940) but not by Brown & Amadon (1968) is a
record from Timor of a specimen of B. Uventer obtained by S. Miiller some-
time between 1826 and 1 8 3 7. It is an old mounted bird in the Leiden Museum
and was listed by Schlegel (1862, Mus. Pays Bas, 2, mon. <f, Buteones: 21-22)
and mentioned by Meyer & Wigglesworth (1898, Birds of Celebes: 49-51).
There is no record otherwise from either Timor or any other of the Lesser
Sunda Islands of any Butastur. As Miiller collected in south west Celebes and
actually obtained B. Uventer there, there is a possibility of an error in labelling
to account for the Timor record, but Dr. Mees informs me that Miiller's
localities are usually reliable. Hellmayr (19 14, Die Avifauna von Timor) and
99
Mayr (1944, Bull. Amer. Mus. Nat. Hist., 83, art. 2) both ignore the Timor
record. As B. liventer has a peculiarly fragmented distribution, the old record
from Timor should not be lost sight of although it needs confirmation.
The Timor record seems inadvertently to have been the basis of citing that
island in the winter range of the Grey-faced Buzzard Eagle, B. indicus, by
Vaurie (1965, The Birds of the Palearctic Fauna — Non Passeriformes : 180).
This species winters in islands east of Wallace's Line but only in the north
(Talaut, Sangihe, Siao, north Celebes, Morotai, Halmahera, Ternate and
Western Papuan Islands).
The Steppe Buzzard, Buteo buteo vulpinus, has once been reported from north
Celebes, for Meyer & Wiglesworth (1898 : 5 1) under the name B. desertorum
state that there is a specimen in the Leiden Museum so labelled and collected
by von Faber in 1883. Stresemann in listing the migrants recorded from
Celebes (1941 ,]ourn. Ornith.: 69-100) ignores the record, either through over-
looking it or because he thought it unreliable. Von Faber did collect in
northern Celebes and Mees (1971, Zool. Meded., 45 : 228) has reported speci-
mens of B. b. vulpinus in the Leiden Museum from Perak, Malaya and from
Java, so that a record from Celebes is not impossible. But Dr. Mees kindly
informs me that there is no such specimen from Celebes in the Leiden
Museum nor do the records provide any evidence that there ever was one.
The statement by Meyer & Wiglesworth cannot be accepted.
The classification of
Tristram's Storm-petrel Oceanodroma tristrami Salvin
by W. R. P. Bourne
Received 16th April, 19J4
While WagstafTe (1972) performed a useful service in locating and confirming
the identity of the type of Oceanodroma tristrami, it should perhaps be pointed
out that he seems to have overlooked a review of this form and its allies by
Austin (1952). The latter points out that it is a member of a group of four
very similar large, dark, fork-tailed storm-petrels which replace each other
in different parts of the Pacific, in which the two occurring over warm water
have generally brownish plumage and the two occurring over cooler water
are bluer. He considers that since each shows some distinct character they
should all be treated as distinct species, including in addition to Tristram's
Storm-petrel Oceanodroma tristrami (pale wing bar, north-west Pacific),
Matsudaira's Storm-petrel O. matsudairae (pale primary shafts, west tropical
Pacific), the Black Storm-petrel O. melania (long tarsus, east tropical Pacific)
and Markham's Storm-petrel O. markhami (short tarsus, south-east Pacific).
I have also suggested (in Palmer, 1962) that the extinct large white-rumped
Guadalupe Storm-petrel Oceanodroma macrodactjla of the north-east Pacific
may also have belonged to the same group, showing a greater development
of the tendency to paleness of the rump also seen in O. tristrami, although of
course like other members of the group it could well have evolved inde-
pendantly from the main body of smaller storm-petrels.
References :
Austin, O. L., Jr. 1952. Notes on some petrels of the North Pacific. Bull. Mus. Comp.
Zool. Harvard 107 ■: 390-407.
Palmer, R. S. 1962. Handbook of North American Birds \ Vol. 1, Loons through Flamingoes.
Yale University Press, New Haven and London.
WagstafTe, R. 1972. The status oiOceandroma tristrami Salvin. Bull. Brit. Orn. Club 92 : 27-28.
Rufous-bellied Heron in Kenya
by Laurence C. Binford <& Dale A. Zimmerman
Received )rd Aprils 1974
On 15 th October 1973, we observed and photographed a Rufous-bellied
Heron Butorides rufiventris (formerly Erythrocnus; see Smithers et al. 1958) at
the edge of Loginya Swamp in the Amboseli Masai Game Reserve, approxi-
mately 2 • 5 km southeast of the new Amboseli Lodge along the Ol Tukai-
Kilaguni road. Colour transparencies (on file in the National Museum,
Nairobi) were obtained with a 450 mm lens and also through a tripod-
mounted, X 20 telescope. Although of rather poor quality, these photographs
are clearly representative of the species. Despite the misleading map in
Mackworth-Praed & Grant (195 2), B. rufiventris apparently has not heretofore
been reported from Kenya. Williams (1964) and Forbes- Watson (1971)
include only Uganda and Tanzania within its East African range.
The heron was feeding in shallow water between 70 and 80 metres from
the road, and we observed it for some thirty minutes with the telescope and
with 8x50 and 8 x 40 binoculars. Twice during this period it flew for very
short distances, settling again into the shallow water. Feeding nearby were
Squacco Herons Ardeola ra//oides, Black-winged Stilts Himantopus himantopus,
Long-toed Lapwings Hemiparra crassirostris, several sandpiper species, and
some Yellow Wagtails Motacilla flara.
The Rufous-bellied Heron appeared to be slightly smaller than a Squacco
Heron and larger, stockier, thicker-necked, and heavier-billed than a Green-
backed Heron Butorides striatus. Its outline was unique and readily distin-
guishable from that of the commoner heron species. The crown and sides of
the head were recorded by us as medium bluish-grey, merging to darker
slate-grey on the neck and upper back. The lower back was dark brownish-
grey. The throat and a continuing short streak down the upper neck were
whitish. The tail, upper tail covers, belly, and at least part of the crissum
were conspicuously rufous, the tail colour showing especially well in flight.
There was considerable rufous on the wings as well.
Soft part colours in this species vary. Jackson (1938) wrote of the adult,
"Iris bright yellow, bare skin round the eye yellow; bill black, basal half
yellowish- white; feet dusky yellow". Chapin (1932) recorded a female with
" . . . orbit greenish yellow; beak black above, changing to greenish yellow
on the lower part; feet lemon yellow". Our bird exhibited the following
colours: a narrow line (presumably bare skin) around the top and rear of the
eye white; bare lores pale yellowish-white; irides pale yellow; bill blackish
with basal two-thirds of the lower mandible dusky orange-yellow; feet bright
orange-yellow.
After leaving the site of this observation and motoring a few hundred
metres along the marsh edge, we encountered Mr. W. G. Harvey, who
showed Zimmerman a second Rufous-bellied Heron. It was similarly
plumaged to the one described above but possessed a more obviously
streaked foreneck and perhaps a slightly paler dorsum.
B. rufiventris appears to be uncommon, or rare everywhere in East Africa,
where the majority of records are from Lake Victoria. The Amboseli Masai
area, with its marshy expanses, would seem a logical place for wandering
or migrating individuals to occur. Possibly they have been overlooked
heretofore.
References :
Chapin, J. P. 1932. The birds of the Belgian Congo Part I. Bull. Amer. Mus. Nat. Hist. 65 :
756 pp.
Forbes- Watson, A. D. 1971. Skeleton checklist of East African birds. Nairobi: Nat. Mus.
(cyclostyled).
Jackson, F. C. 1938. The birds of Kenya Colony and the Uganda Protectorate 1. London: Gurney
& Jackson.
Mackworth-Praed, C. W. & Grant, C. H. B. 1952. African handbook of birds. Series 1, Vol. 1.
London: Longmans.
Smithers, R. H. N. et. al. 1958. Second report of the S.A.O.S. list committee. Ostrich 29:
23-46.
Williams, J. G. 1964. A field guide to the birds of East and Central Africa. Boston: Houghton
Mifflin.
Geographical variation in iris colour in the
bulbul Andropadus milanjensis
by R. J. Dowse tt
Received 1 jth February \ 1974
The Stripe-cheeked Bulbul Andropadus milanjensis (Shelley) is a species of the
montane forests of eastern Africa. It is apparently most closely related to
Andropadus tephrolaemus (Gray), but in the few localities where the two are
geographically sympatric milanjensis occurs at a lower altitude than teph-
rolaemus (C. W. Benson in lift.). Moreover milanjensis may make some seasonal
movements to lower levels still. In August Benson (1937) found it numerous
in riparian scrub at 1,370 m in the Misuku Hills in northern Malawi, but not
at all in the larger areas of forest at 1,830 m. Yet in October I found this
species in breeding condition and common in the Misuku forests at 1,830 m
and above. Specimens from Mlanje in southern Malawi were collected by
Benson (in litt.) from July to September from as low as 760 to 1,220 m. A.
tephrolaemus is less confined to the forest interior than is milanjensis ', although
the latter is at times found in very thin riparian growth. A. tephrolaemus is
often on the edge of high altitude forest, even in isolated trees in adjacent
grassland. The relationship between these two species is discussed by Hall &
Moreau (1970).
Three subspecies of A. milanjensis are recognised by White (1962), and in
addition Clancey (1969) has described disjunctus from eastern Rhodesia and
adjacent Mocambique. Most of the Tanganyika-Nyasa montane group is
occupied by olivaceiceps (Shelley) (type locality: Chiradzulu, southern Malawi).
This race ranges from Cholo in southern Malawi east of the Shire rift, north-
westwards to the Rungwe area of south-western Tanzania. It is replaced by
striifacies (Reichenow & Neumann) (type locality : Marangu in north-eastern
Tanzania) from the Iringa area, northwards. Within its range olivaceiceps is
local in distribution, being absent from several montane forests for no
known reason (see Benson 1953). In north-central Malawi, between Nchisi
and the eastern escarpment of the Nyika Plateau, there is but one record, a
specimen collected at Mwanjati on the southern Vipya Plateau (Benson 195 1).
Although Benson (1953) did not indicate it, he (in litt.) believes the species
to be absent from the Vipya (see also Benson 1941, Ibis (14)5 : 5 5-Ed.), which
he knows well, except in the extreme south. It is apparently absent from
Dedza, Chongoni and Dzalanyama in central Malawi, although present on
the equally small Chirobwe Mountain (Benson ibid).
Within the subspecies olivaceiceps the colour of the iris of adults appears to
be either yellowish (pale) or reddish (dark). Adults collected from Cholo in
southern Malawi north to Nchisi in central Malawi all have irides described
by the collectors as a shade of yellow or yellowish-white. On the other hand
adults from the Nyika Plateau and Misuku Hills in northern Malawi all have
irides a shade of umber or chestnut. The Table summarises the iris colour
TABLE
Iris colour in adult Andropadus milanjensis olivaceiceps
Area Colour Sample
Misuku Hills
pale salmon, brick
56*6% i?
Nyika Plateau
pale reddish brown
i(J
Nchisi Mt.
yellowish, pale yellow
16", i?
Chirobwe Mt.
pale yellow
i?
Mangochi Mt.
yellowish-white, pale yellow
io6U i?
Zomba Plateau
pale yellow
i?
Chiradzulu Mt.
pale yellow
16*
Soche Mt.
pale yellow
io*
Cholo Mt.
naples yellow, yellowish brown
*<3<3
recorded for adult specimens in the British Museum at Tring (kindly
examined for me by C. W. Benson) and of recent specimens collected by me
for the Livingstone Museum in Zambia. It will be seen that the differences
demonstrated cannot be sexual; neither can they be seasonal, as shown by
series collected by me in October from Mangochi Mountain (io, yellow
irides) and the Misuku Hills (5, near brown irides).
Specimens from northern Mozambique (Namuli Mountain) and eastern
Rhodesia (Chirinda, Mt Selinda) have brownish irides. Both populations
are attached to the nominate race by White (1962), but the Rhodesian birds
are what Clancey (1969) has called disjunctus. More than 30 disjunctus collected
by Mr. P. A. Clancey all had umber brown irides (R. K. Brooke in ////.). The
iris colour of the type specimen of A. milanjensis from Mlanje was not
recorded by the collector, but two adults and a juvenile from the same area,
collected by Benson, have irides pale grey. Striifacies from the following
localities also have the iris recorded as grey on the labels of specimens in the
British Museum: Chyulu Hills, south-eastern Kenya; Moshi, Kilimanjaro;
Monduli Mt., northern Tanzania. On the other hand one specimen from this
last locality has the iris labelled "brown", and one from Kilomeni in the
North Pare Mts. as "yellow". Hall & Moreau (1970) refer to northern birds
as having yellow or pale eyes. Unfortunately further specimens labelled with
iris colour are not available, and the situation in the northern populations
(i.e. striifacies and olivaceiceps north of the Misuku Hills) requires further
investigation. Juvenile birds from eastern Rhodesia (Vumba 1) and northern
Malawi (Mwanjati, southern Vipya 1 ; Nyika 1) have irides respectively
greyish-brown, brownish-yellow and pale grey-brown. The iris colour of
adults at Mwanjati on the southern Vipya is not known, but this area is
between the known extremes of brown (Nyika) and yellow (Nchisi).
There are no apparent differences in plumage colours between these two
groups of olivaceiceps. There may be a difference in average size, as suggested
by the following weights and measurements of my own Mangochi and
Misuku samples, but further data are required to confirm this : —
n
Wing (mm)
Tail (mm)
Weight (g)
Misuku Hills 4
97-100 (99-0)
92-94 (93-0)
40-9-46-5 (44-0)
Mangochi Mt. 7
95-101 (97-7)
85-92(89-3)
37-0-43-5 (40-0)
103
These data are of adult males only, all of which had enlarged testes.
The existence of two, apparently consistent, iris colour groups within the
subspecies olivaceiceps suggests that these populations are genetically different.
Although these differences cannot be noticed in preserved skins they are
likely to have an important biological basis. A specimen from the type
locality of olivaceiceps (Chiradzulu) has yellowish irides, and therefore the
northern populations of brown-eyed birds may need to be recognised as
subspecifically distinct. But I do not intend at this stage to present a new
subspecific name. Clearly a detailed study of iris colour throughout the range
of A. milanjensis will be necessary to determine the significance of the ob-
served differences.
Although the Shire rift in southern Malawi restricts the distribution of
several montane birds, to the extent that Dowsett (1971) has separated the
mountains east of the rift into a South-eastern montane group, this is not
the boundary between the two populations of olivaceiceps. However, the
apparent gap between the two groups occurs at another natural geological
break: Nchisi Mt. (i3°2 5'S.) and the southern Vipya Plateau (i2°4o'S.) are
separated by some no km of low country, mostly below 910 m altitude. In
particular the valleys of the Bua and Dwangwa Rivers are likely to have
formed effective barriers to montane bird dispersal for a long period in recent
time.
Acknowledgements : I am most grateful to C. W. Benson for examining
specimens at the British Museum, Tring, on my behalf, and to the staff there
for their usual kind co-operation. My thanks go to R. K. Brooke, P. A.
Clancey and M. P. Stuart Irwin for useful comments on this problem.
References :
Benson, C. W. 1937. Miscellaneous notes on Nyasaland birds. Ibis (14)1: 551-582.
— 195 1. Breeding and other notes from Nyasaland and the Lundazi District of Northern
Rhodesia. Bull. Mus. Comp. Zool. Harv. 106: 69-114.
— 1953. A check list of the birds of Nyasaland. Blantyre & Lusaka: Joint Publications
Bureau.
Clancey, P. A. 1969. Miscellaneous taxonomic notes on African birds, 27. Durban Mus.
Novit. 8 (15): 227-274.
Dowsett, R. J. 1 97 1. The avifauna of the Makutu Plateau, Zambia. Rev. Zool. Bot. Afr.
84 (3/4): 3 1 2 -3 33-
Hall, B. P. & Moreau, R. E. 1970. An atlas of speciation in African passerine birds. London:
Brit. Mus. (Nat. Hist.).
White, C. M. N. 1962. A revised check list of African shrikes, orioles, drongos etc. Lusaka: Govt.
Printer.
The status of the cormorants
Phalacrocorax carbo lucidus and Phalacrocorax carbo patricki
by Emil K. Urban <& T. G. Jefford
Received nth February, 1974
Cormorants are common along the Kazinga Channel in Uganda's Ruwen-
zori National Park. Some adults there have black underparts except for white
which extends from the cheeks and throat along the entire front of the neck
to the upper breast; this description corresponds to that of P. c. lucidus
(Lichtenstein), the White-necked Cormorant of sub-Saharan Africa. Also
occurring along the Kazinga Channel are individuals of P. carbo which have
the white restricted to the throat while the neck and upper breast are black
104
and still others whose feathered parts are wholly black. Black-necked
individuals with white throats resembling P. c. sinensis (Shaw) were de-
scribed by Williams (1966) as a distinct subspecies of carbo, P. c. patricki. He
suggested that the white-necked form of sub-Saharan Africa should therefore
be known as "Pbalacrocorax lucidus lugubris Ruppell, not P. carbo lugubris"
(=P. c. lucidus \n White 1965).
Williams's observations therefore indicate that in the Kazinga Channel
there are two distinct sympatric species of large cormorants — one with a
white neck, the other with a black neck. Our observations, however,
show that there are intermediates of all types between the forms described
by Williams as patricki and lugubris \ some of these forms are depicted in
Figure 1 and numbers of birds assigned to four categories based on the extent
of white plumage from the head towards the belly are given in Table 1.
\ b c c d
Figure 1. Several cormorants drawn from 35 mm slides taken at Kazinga Channel, show-
ing variation in throat and neck plumage from all black to white extending from throat
along neck onto upper breast. Individuals A, B, C & D are referred to in Table 1 .
Table i
Counts of the plumage types of cormorants from the Kazinga Channel, Uganda. Birds with
black bellies were assigned to the appropriate categories, with the results given in this
table. Representatives of these categories are shown in Figure 1. Birds with white bellies
are immature, and were not included. Category: A, no white on throat and neck; B, throat
and less than \ neck white; C, throat and h to all neck white; and D, white extending be-
yond neck onto upper breast.
Category A B C D
Number of individuals 27 29 45 18
% of total counted
15-1
Williams's assignation of the black-necked birds to P. c. patricki and the
white-necked birds to P. lucidus lugubris was made because he saw no cor-
morants with intermediate plumages and because the black-necked birds
which he collected were in breeding condition. It would be reasonable to
assume therefore that the patricki birds breed with one another, rather than
with lucidus birds. However, M. P. L. Fogden (pers. comm.) has since
observed that at a nesting colony at Lake Murabyo (o°o5'S, 3o°o2'E) within
105
the Maramagambo Forest in the Ruwenzori National Park, the black-necked
and white-necked birds pair without any apparent regard to the amount of
white on the neck of the partner.
Cormorants with various amounts of black and white plumage on the
neck have been reported from other parts of eastern Africa. Chapin (1932)
reported that some specimens of P. carbo from eastern Zaire and Lake
Victoria (Bukoba, Tanzania) have white more restricted to the upper fore-
neck than does typical lucidus. Reichenow (1892a, 1892b) described a specimen
from Bukoba as P. gutturalis, now synonymous with P. carbo (Sharpe &
Ogilvie-Grant 1898; Peters 193 1), whose neck colour is white mixed with
black which extends only to the black upper breast. On 3 1 July 1970 we saw a
nesting adult at Lake Abiata, Ethiopia, whose throat, cheeks and upper third
of the neck were white while the lower two-thirds of the neck were black
necked with a few white spots. A. Kortlandt (pers. comm.) has seen P. carbo
at a breeding colony near Bukavu on Lake Kivu (Zaire) with neck colours
either black or white. Lippens (1938) reported P. carbo at Lake Edward
(now Idi Amin Dada) in Zaire, with neck colours varying from all white to
all black, with white in the latter restricted to the throat. At Ishango, where
the Semliki River drains from Lake Edward, both black-necked and white-
necked forms are present, but the former are in a pronounced majority (K.
Curry-Lindahl pers. comm.; see also Curry-Lindahl 1961: 146-149).
Thus, the cheek, throat and neck plumage of P. carbo in parts of eastern
Africa may vary from individual to individual, ranging from all black to all
white. Apart from the estimate given in Table 1 , we are not aware of any
figures from which the proportions of the various forms in different popula-
tions can be estimated nor if proportions within the populations are in
equilibrium or are changing. However, it may be stated that the black-
necked form is centred in western Uganda and eastern Zaire, although the
white-necked form always seems to be present as well, and that the amount of
black in forms with black and white becomes progressively less the farther
individuals are found from this part of Africa.
We suggest that the black/white-necked cormorants in eastern Africa
represent variations in plumage within a single species, P. carbo {lucidus), and
not two species, P. lucidus (lugubris) and P. carbo (patricki), as proposed by
Williams (1966). We do so because of the following reasons: (1) we know of
no evidence of any "pure" breeding populations of patricki\ (2) where
black-necked and white-necked forms occur together, there is a whole
series of variants from those with all black necks through various inter-
mediate stages to all white necks; (3) in entirely white-necked forms, even
when there are no black-necked forms present, there is variation in the
amount of white which extends toward the black belly; (4) rare individuals
with partly black necks occur in breeding populations of P. carbo lucidus as
far away as Ethiopia; and (5) variation in plumage within a species is not
uncommon in the genus Phalacrocorax. For example, in New Zealand
Phalacrocorax melanoleucos (Vieillot) 1 has mixed forms at the breeding colony,
varying from birds with a black under surface to those with a white one
(Oliver 1955). A similar situation occurs in P. carunculatus chalconotus (G. R.
Gray) 1 in that breeding populations in New Zealand and associated islands
have 'two distinct colour phases' (Oliver 195 5). The variation in plumage of
the New Zealand and the eastern African cormorants indicates that at times
it is no simple matter to separate species within the genus Phalacrocorax.
1 See footnote at end of text.
106
The nature of the selective pressures which might establish and affect these
variants in eastern African cormorants is obscure. Interesting questions arise.
For example, a variety of birds which prey on aquatic animals, e.g. fulmars,
boobies, herons, egrets and skuas, exhibit light and dark phases; the main-
tenance of these different forms has been ascribed to different efficiency in
different feeding situations (Simmons 1972). Is this also a factor in the case of
black/white-necked cormorants of eastern Africa ? Also, why are the black-
necked forms concentrated largely in western Uganda and eastern Zaire ? Is
this darker stock a relic of colder, wetter periods in the past when carbo in
Africa may have been like the black-necked populations of carbo in the
Palearctic of today? Furthermore, why do most carbo in sub-Saharan Africa
today have white necks? Did possession of white plumage on the neck
evolve largely as an advantageous response to heat stress at the nesting
colonies as our studies in Ethiopia suggest (Jefford & Urban in prep.) ?
It is clear that the resolution of these questions requires much more
detailed data on the distribution of the various forms and on their juvenile
and breeding plumages, and the correlation of these data with environmental
factors and the biology of the birds. It seems equally clear to us that based
on such information as is now available, there is no justification for removing
lucidus from the species P. carbo nor for describing the black-necked form as
the subspecies patricki.
We acknowledge with thanks the financial support of African Wildlife
Leadership Foundation, Haile Sellassie I University, National Geographic
Society and University of Miami's Department of Biology and Maytag
Chair of Ornithology. We are most grateful to the Chapman Fund of the
American Museum of Natural History for financial support which enabled
the second author to observe cormorants in the Kazinga Channel and sur-
rounding areas, to the National Research Council of Uganda for permission
to do so, and to Dr. S. K. Eltringham, Dr. D. E. Pomeroy, and many mem-
bers of the Uganda National Parks for much assistance in Uganda.
1 We follow Peters's (1931) specific and subspecific classification in which the species
melanoleucos is divided into two subspecies, melanolencos melanoleucos and melanoleucos
brevirostris (Gould). Oliver (1955) however lists melanoleucos and brevirostris as separate
species or in his words 'semispecies'. Also he separates Peters's (193 1) P. carunailatus
chalconotits into two forms or 'semispecies', P. huttoni and P. cbalconotus.
References :
Chapin, J. P. 1932. The birds of the Belgian Congo. Part I. Bull. Amer. Mus. Nat. Hist.
65: 756 pp.
Curry-Lindahl, K. 1961. Contribution a r etude des vertebres terrestres en Afrique tropicale.
Exploration du Pare National Albert et du Pare National de la Kagera. Fasc. 1. Inst. Pares
Nat. du Congo Beige et du Ruanda-Urundi.
Lippens, L. 1938. Les oiseaux aquatiques du Kivu. Le Gerfaut 28: 1-196.
Oliver, W. R. B. 1955. New Zealand birds, 2nd ed. Wellington, N.Z.: A. H. & A. W. Reed.
Peters, J. L. 193 1. Cheek-list 0/ birds of the world. Cambridge, Mass.: Harvard University Press.
Reichenow, A. 1892a. Zur Vogelfauna des Victoria Njansa. Journ. Ornith. 40: 1-60.
— 1892b. Bericht liber die Dezember-Sitzung. Journ. Ornith. 40: 129-134.
Sharps, R. B. & Ogilvie-Grant, W. R. 1898. Catalogue of the birds in the British Museum 26.
London: Brit. Mus. (Nat. Hist.).
Simmons, K. E. L. 1972. Some adaptive features of seabird plumage types. Brit. Birds 65 :
465-479, 510-521.
White, C. M. N. 1965. A revised cheek list of African non-passerine birds. Lusaka: Govern-
ment Printer.
Williams, J. G. 1966. A new cormorant from Uganda. Bull. Brit. Orn. CI. 86: 48-50.
107
Relative Biomass of Ethiopian and
Palaearctic Passerines in West Kenya Habitats
by Peter L. Br it ton
Received ph February, 1974
The problem of the accommodation in Africa of vast numbers of migrants
from the Palearctic region has attracted considerable attention during the last
two decades, mainly through work and stimulus of the late R. E. Moreau.
The various carefully considered estimates given by Moreau (1972) for the
total mean number of migrants supported by sub-Saharan Africa range from
1600 millions to 3750 millions. Impressive though these figures are, they
represent only one or two birds per hectare in the approximately 20 million
km 2 involved, assuming a uniform distribution throughout the reception
area. It is now well known that African evergreen forest (except the edges)
and montane areas are mainly avoided by migrants, so that favoured habitats
would be expected to contain a far greater density; and the geographical
distribution of migrants in Africa is far from uniform, with the northern
tropics receiving far more species than areas further south. Pearson (1972)
found as many as 75 wintering Palearctic passerines per hectare in lake side
vegetation near Kampala, southern Uganda, but as few as 2 - 5 birds per
hectare in cultivated bush away from the lake. Morel (1968) estimated only
one or two wintering Palearctic passerines per hectare in Senegal thornbush.
Moreau (1972) has shown that about 20 per cent of the African non-forest
avifauna consists of immigrant species. Bearing in mind that there are
virtually no granivorous species involved in the immigrant avifauna, the
figure is increased, for example to 26 per cent of the non-granivorous, non-
forest avifauna in the northern tropics. The data in Morel (1968), for Senegal,
yield a still higher figure, with 3 5 per cent of all species in his study area of
Palaearctic origin. Such an analysis by species is far from satisfactory in
assessing the impact of the immigrant avifauna, and Moreau had to admit
that "of the comparative numbers of individuals we know nothing". And
referring to the figure of 26 per cent above: "with no knowledge of the
number of individuals involved it would of course not be justifiable to
conclude that in this part of Africa one quarter of the food available other
than grain is diverted to the migrants".
As residents of western Kenya from September 1968 to December 1972,
my wife and I netted and ringed large numbers of birds in various west
Kenya habitats. Some of these data permit valid estimates to be made of the
relative numbers of Ethiopian and Palaearctic passerines. Data from netting
are preferred to visual counts, since estimates are more objective and avoid
the conspicuousness factor. Naturally, some species avoid capture more
readily than others, and species which normally forage more than about 3 m
from the ground are seldom captured. In forest habitats it is evident that only
forest undergrowth birds are sampled using this technique, but this is not a
serious problem in the other habitats discussed, there being comparatively
little vegetation above net height. A large number of Palaearctic passerines
are ringed annually in East Africa, but most are captured in such a way that
no useful comparisons with Ethiopian species can be made. In the main, sites
known to be especially prolific in migrant species at a particular time are
worked, but there is virtually no netting between May and August. For valid
comparisons there must be no such Palaearctic bias.
Between September 1971 and November 1972, a census was conducted in
108
four habitats in Siaya District, Central Nyanza, mainly using mist-nets, but
also counting birds present in the study plot. One visit a month over the
fifteen-month period was made to each plot. The same nets were set in the
same sites within each plot on each visit, and were set for the first four hours
of daylight each time. All mornings were calm, and dry so that any variation
in totals captured as a result of varying weather conditions was minimal.
With nets set only once in a month, net shyness did not apparently reduce
totals (see Table i). Three of the plots were one hectare in size, but the more
open acacia area was two hectares. The quantity of netting used in each area
was the same (300ft., ca.cjom). Table 1 shows monthly passerine captures;
Table 2 shows the totals of species recorded during the study period,
visually or otherwise; Table 3 shows passerine captures in October-Novem-
ber, when Palaearctic migrants were on southward passage; and Table 4
shows winter data, for December-January only, including data from forest
areas (see below). Granivorous species (taken to include all Fringillidae,
Estrildidae, Ploceidae) are excluded from Tables 3 and 4. Biomass data are
based on individual weights of all birds caught, recorded on Pesola spring
balances. Drawing on the census results two longer papers are in preparation
on detailed aspects of the papyrus and savanna avifaunas.
It is evident from Tables 1-4 that, of the four habitats worked, the thicket
and acacia habitats are the most important for Palaearctic migrants. But even
here, counts or estimates of Palaearctic passerines per hectare, aerial feeders
excluded, never exceeded 30 birds at times of passage, 15 birds in winter.
Large stands of Cyperus papryrus form a remarkably homogenous habitat
with a highly specialized resident avifauna and an extremely low avian
species diversity index (Britton in prep.). Palaearctic species feed along the
edges but hardly penetrate the interior. The six birds netted in papyrus in
December 1971 were feeding in an area which had been cleared and planted
with cabbages during an exceptionally dry period. These figures are atypical
and are excluded from Table 4, in which only three of the study plots are
included. The habitat termed cultivation in Tables 1-4 is a heterogeneous
habitat, which includes overgrown farmland, remnant thicket and streamside
vegetation. It is especially rich in granivorous species. The two habitats
favoured by Palaearctic species are:
(a) thicket, mainly secondary growth but impenetrable, including about
30 per cent introduced Lantana camara, intermixed with indigenous vege-
tation; altitude 1450 m. Eighty per cent of vegetation is 3-4 m high with
only occasional trees as high as 6 m. Apart from the Lantana, dominants are
Vangueria acuti/oba, Combretum molle, C. binderianum, Albi-^ja coriaria, Bridelia
micrantha, Vitex doniana. V. fischeri, and others.
(b) well-grazed, seasonally inundated Acacia seyal savanna, altitude 1 200 m.
Trees up to 4- 5 m but seldom taller than 2 • 5 m, with a density ranging from
about 35 to 65 trees per 1000 m 2 .
Moreau (1970) and Pearson (1972) have mentioned the importance of
introduced Lantana berries as a food for Palaearctic migrants in Africa, in
particular for the Garden Warbler Sylvia borin. During October and Novem-
ber, 60 per cent of non-granivorous birds (50 per cent of biomass) in the
thicket plot were of Palaearctic origin (see Table 3). Virtually all were S.
borin and their faeces contained little else except Lantana berries. In December
and January the percentage of Palaearctic birds was reduced to 27 per cent
(25 per cent of biomass), also mainly S. borin (Table 4). December-January
figures for other habitats, including forest undergrowth, range from 2 to 19
per cent of individuals, 1 to 1 1 per cent of biomass (Table 4).
109
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Acacia sejal savanna is an important habitat for passage migrants but it
provides regular winter quarters for only the Whinchat Saxicola rubetra (one
or two per hectare). October-November figures for this habitat in Table 3
are inflated by a large catch of aerial feeding Swallows Hirundo rustica and
Sand Martins Riparia riparia, so that Palaearctic species account for 52 per
cent of non-granivorous birds (44 per cent of biomass). Concentrations of
feeding hirundines are not normally found in this plot, but even in their
absence the percentage of Palaearctic birds caught can be large when a "fall"
of migrants is involved. For example, morning catches were made in this
habitat, but outside the study plot, on 9th November 1969 and 22nd March
1 97 1 : of 27 non-granivorous passerines caught on the first date, 16 were of
Palaearctic origin (1 1 species), forming 59 per cent of the total (46 per cent of
biomass); and of 9 non-granivorous passerines caught on the last date, 7 were
of Palaearctic origin (5 species), forming 78 per cent of the total (83 per cent
of biomass). These figures contrast with only 4 per cent of the October-
November catch in papyrus only 1 km away (Table 3). Yet, considering
species rather than individuals, the papyrus plot has the highest percentage
of Palaearctic birds (Table 2).
Data from other habitats were collected in a less systematic manner,
without any special effort to catch Palaearctic birds. Table 4 includes data
from Kakamega Forest, 50 km east of the study area, and from three sites on
Mt. Elgon, 150 km north of the study area. Nets at Kakamega were set along
forest paths and captures do not include any edge element, but Elgon sites
TABLE 2
Totals of passerine species recorded in four Central Nyanza study plots in fifteen monthly
Palaearctic
species as %
of non-
Non-granivorous
Palaearctic
granivorous
All
species
species
species
species
Thicket (area 1 ha)
48
39
9
23
Acacia (2 ha)
64
43
11
26
Cultivation (1 ha)
71
42
8
19
Papyrus (1 ha)
48
30
9
30
TABLE 3
Palaearctic passerine captures on four Central Nyanza study plots in October-November
Thicket Acacia Cultivation Papyrus Total
Hirundo rustica
15
15
Riparia riparia
8
8
Motacilla flava
1
1
Muscicapa striata
1
1
Ficedula albicollis
1
1
Laniiis cristatus
3
3
Saxicola rubetra
4
4
Sylvia borin
2 7
2
8
1
38
S. atricapilla
1
1
Acrocephalus scirpaceus
1
1
2
Phylloscopus trochilus
1
1
Total of Palaearctic birds
29
34
10
2
75
Total of non-granivorous birds
48
66
39
50
Biomass (g) of Palaearctic birds
535
578
178
30
Biomass (g) of non-granivorous
birds
1062
1291
903
829
Palaearctic birds as % of total
60
5 2
26
4
Palaearctic biomass as % of total
50
44
in
20
4
were often at the edge. The number of Palaearctic birds at Kakamega (2 per
cent) would certainly increase if edge areas were worked ; but it is doubtful
whether figures would be as high as on Mt. Elgon at much higher altitudes
(10-12 per cent). Forest undergrowth above about 1500 m provides winter
quarters for only the Blackcap Sylvia atricapilla, which accounts for virtually
the whole Palaearctic catch in these areas.
Lake Nakuru in Kenya's Rift Valley, some 150 km south-east of the study
area, provides a further comparison. The data in Table 5 refer to December
1968 when the lake level was high. Nets were set for thirteen days at the lake
edge, in sedges and grassland with scattered small acacia trees, and were
hardly closed except at night. Hirundines formed 8 1 per cent of the total
passerine catch, and two species of Palaearctic hirundines alone formed 49
TABLE 4
West Kenya habitats as winter quarters for Palaearctic passerines, December-January
data only
Thicket Acacia Cultivation Kakamega
forest
(1700m)
3
Mt. Elgon forest
2400m 2500m zi
8
4
3
21
146
56
57
651
4022
19
2
9
1
10
5
20
100
47
161
156
86
37i
2099
986
3652
10
11
12
7
9
10
Sylvia borin 7
S. atricapilla
A.crocephalus palustris
A., schoenobaenus
Phylloscopus trochilus
Saxicola rubetra 4
Phoenicurus phoenicurus 1
Motacilla flava 1
Total of Palaearctic
birds 8 5
Total of non-
granivorous birds 30 27
Biomass (g) of
Palaearctic birds 149 78
Biomass (g) of non-
granivorous birds 602 729
Palaearctic birds as
% of total 27 18
Palaearctic biomass
as % of total 25 11
TABLE 5
Lake edge at Nakuru as a winter habitat for Palaearctic passerines December 1968 data only.
Palaearctic species No. Biomass (g)
Hirundo rustica 235 4J5 1
Riparia riparia 49 614
Moticilla flava 5 8 947
Phylloscopus trochilus 6 50
Lanius cristatus 3 76
Muscicapa striata 1 14
Acrocephalus schoenobaenus 1 11
Total 353 5863
Ethiopian species
Riparia paludicola 184 1806
Other non-granivorous species 43 959
Total 227 2765
Palaearctic birds as % of total • 60 68
Palaearctic hirundines as % of total 49 55
Riparia paludicola as % of total 32 21
Hirundines as % of total 81 76
Palaearctic hirundines as % of hirundines 61 73
per cent of the catch. Palaearctic passerines as a whole formed 60 per cent of
the catch (68 per cent of biomass). The vast majority of these Palaearctic
birds were either Swallows, Sand Martins or Yellow Wagtails Motacilla flava,
probably mainly wintering birds. As such, these figures are remarkably high,
especially bearing in mind that figures for non-passerines (mainly waders)
would likely show a far greater percentage of Palaearctic individuals.
SUMMARY AND CONCLUSIONS
Moreau (1972) has shown that about 20 per cent of the African non-forest
avifauna consists of Palaearctic immigrant species, but there is no estimate
available of the comparative numbers of individuals. Data obtained from
systematic netting in various west Kenya habitats permit valid estimates to
be made. Only non-granivorous passerines are considered.
During the passage months of October and November, Palaearctic
passerines (mainly Sylvia boriti) constitute 5 o per cent of the avian biomass in
Lantana thickets ; and the percentage (various species) may be still greater in
Acacia seyal savanna, though on average it is much less if aerial feeding
hirundines are excluded. In overgrown cultivation the figure is 20 per cent,
and in papyrus swamp only 4 per cent. Considering species only, papyrus has
the greatest proportion of Palaearctic birds (30 per cent), which illustrates
how misleading figures based on species rather than individuals can be in
assessing the impact of the immigrant avifauna.
As a wintering area, Lantana thickets are again important, with Palaearctic
species accounting for 25 per cent of the avian biomass, against figures of
1 — 1 1 per cent in other habitats, including forest undergrowth. Away from
the edges, forest undergrowth and papyrus swamp provide haven for
virtually no Palaearctic birds. The further inevitable destruction and frag-
mentation of forest and swamp habitats will increase the feeding oppor-
tunities for Palaearctic migrants, and favour the further dissemination of
obnoxious plants like Lantana camara.
A sample from Lake Nakuru in December 1968 is included, taken when
the lake level was high, and assumed to refer to wintering birds. In lake edge
sedges and grassland, Palaearctic species accounted for 68 per cent of the
avian biomass, almost entirely Hirundo rustica, Riparia riparia and Motacilla
flava.
References :
Moreau, R. E. 1970. Changes in Africa as a wintering area for Palaearctic birds. Bird Study
17: 95-104.
— 1972. The Palaearctic- African bird migration systems. London & New York: Academic
Press.
Morel, G. 1968. Contribution a la synecologie des oiseaux du Sahel senegalais. Mem.
O.R.S.T.R.O.M. 29.
Pearson, D. J. 1972. The wintering and migration of Palaearctic passerines at Kampala,
southern Uganda. Ibis 114: 43-60.
On the validity and range of hamprotornis
corruscus mandanus Van Someren, 1921
by P. A. Clancey
Received 21st March, 1974
Van Someren (1921) was the first to demostrate that the glossy starling
Lamprotornis corruscus Nordmann, 1835: Galgenbosch, near Thornhill,
eastern Cape, is polytypic when he proposed L. c. mandanus (van Someren),
113
1 92 1 : Manda Island, coastal Kenya, on the basis of small size. Sclater (1924-
1930) admitted mandanus, giving its southern range limits as the lower
Zambesi R. In 1952, I pointed out (Clancey 1952) that north-eastern Zulu-
land specimens appeared to be inseparable from mandanus from East Africa,
but in my own Catalogue (Clancey 196 5-1 966) I admitted only the nominate
subspecies as occurring in the South African Sub-Region. Mackworth-Praed
& Grant (195 5.) recognise all four described subspecies: L. c. corruscus, L. c.
mandanus, L. c.jombeni and L. c. vaughani, showing mandanus as ranging south
to the littoral of northern, Mozambique. Amadon (1956: 1962) synonymized
with nominate corruscus both mandanus and Van Someren' s later second men-
sural raceZ>. c.jombeni (van Someren), 193 1 : Jombeni Hills, north-east of Mt.
Kenya, Kenya, admitting only corruscus and the insular L.c. vaughani (Banner-
man), 1936: Pemba Island, Tanzania.
During a recent working visit to the National Museum of Rhodesia,
Bulawayo, the Keeper of Ornithology, Mr. M. P. Stuart Irwin, mentioned to
me that Blackbellied Starlings breeding in the Haroni-Lusitu area of south-
eastern Rhodesia were certainly not the same as those of the eastern Cape
(topotypical of L. c. corruscus). I have now investigated this matter with a
reasonable panel of material which reveals that L. c. mandanus is a valid race
with an immense range, extending from the Juba R., in Somalia, south to
Zululand (Ngoye Forest) in South Africa (confirming my tentative findings
of 1952), and that L. c. corruscus is restricted to the south-eastern and eastern
Cape and Natal. The size parameter cannot be employed satisfactorily unless
one has adequate series of both adult and immature (pre-basic) birds and the
sexing of the specimens is beyond reproach. I mention this because much of
TABLE 1
The wing-length variable in adults of three subspecies of
Lamprotornis corruscus Nordmann
Population
Sex
N
Range
M
S.D.
Lamprotornis
corruscus corruscus
Eastern Cape
s
11
113-5-119
115-8
1-85 mm
?
6
108-111
109-2
1-36
Pondoland
6*
9
113 -5-118
115-3
i- 80
?
7
107-5-110
108-3
1-18
Natal
0*
8
111-117-5
115-3
2-75
9
8
107-5-112
108-9
1-74
Lamprotornis corruscus mandanus
Zululand (Ngoye)
s
5
109-113
III -o
1-58 mm
? ,
2
106, 106-5
Sul do Save
o* '
7
1 09-1 1 3
IIO-2
1-98
?
6
104-107
I05-8
1-13
Manica e Sofala
3
6
107- 5-112
IIO-O
2-04
?
1
105
Eastern Rhodesia
6*
?
0*
9
6*
8
109-5-112
110-5
0-82
Tanzania
4
108-112
IIO-2
1-71
Kenya (coast and Tana R.)
12
107-5-113
I0 9 -5
i-55
?
10
102-106
I04-I
1-70
Lamprotornis corruscus jombeni
Kenya (highland) (after
6*
5
116-121
II9-0
2-74 mm
Van Someren 1932)
The extremely conservative nature of the wing-length ranges and close similarity of the
means in L. c. corruscus and L. c. mandanus will be noticed.
114
Amadon's material was unsexed and he clearly did not differentiate between
adults and those in immature plumage when formulating his conclusions. In
this starling immature (pre-basic) males are smaller than adults, equalling in
size adult females, while immatures of the latter are substantially smaller than
their respective adults. In arriving at my conclusions I have taken care to sort
out the material into age classes.
Specimens of adults in immaculate plumage from the Juba R. and the coast
of Kenya, south as far as the Ngoye Forest, Mtunzini district, Zululand,
show no differences of subspecific import. Males of this assemblage of popu-
lations have the flattened wings ioy 5-1 13. females 102-107 mm (see Table
1), the males with the ventral surface below the Chessylite Blue (Ridgway
191 2, pi. xx) plastron medially washed with metallic violet and heavily burn-
ished with antique gold sheen. South of Zululand, in Natal, the Transkeian
Territories and the south-eastern and eastern Cape, size in adults increases
markedly, thus: wings of ^ (m) II 3'5 _II 9> $9 108-112 mm. The male
with a wing as low as 1 1 1 mm. is from the Natal population, and reveals the
influence of the presence of the small eastern littoral mandanus in adjacent
Zululand. Ventrally the large-sized southern males show extensive matt
black over the medio-ventral plane, with reduced metallic violet overlay, the
gold sheen restricted to the lateral surfaces and flanks. Females show no
obvious colour differences not consonant with wear or individual variation.
Despite opinion to the contrary, L. c. mandanus is a perfectly valid subspecies,
but is not restricted to East Africa, and requires to have its established range
extended south to Zululand in the Republic of South Africa.
The littoral populations of the Blackbellied Starling ranging from south-
western Somalia to the south-eastern Cape may therefore be arranged in two
subspecies on the basis of the following summarized characters :
(a) Lamprotornis corruscus corruscus
$<$ wings in adults 113-5-119, 9? 108-112 mm. Male with venter below
plastron medially matt black with some violet gloss and washed laterally
with antique gold sheen.
Ranges from the south-eastern and eastern Cape to Natal.
(b) Lamprotornis corruscus mandanus
o*o* wings in adults 107- 5-1 13, 9? 102-107 mm. Male with venter below
the plastron medially overlaid with metallic violet and heavily burnished with
gold sheen. Ranges from south-western Somalia (Juba R.) and Kenya on the
Tana R. and coast, to the mainland coast, Zanzibar and Mafia in Tanzania,
Mozambique, eastern Rhodesia, eastern Swaziland (Lebombo Mts.) and
Zululand.
I have not seen any of the limited available material of L. c.jombeni, but
judging by the wing-lengths of adult males given by van Someren (1932) it
is even larger on average than the nominate subspecies, and is stated by its
describer to be "rather brighter" in colour facies than L. c. mandanus. Its
range is restricted to the eastern highlands of Kenya on Mt. Kenya and the
neighbouring Jombeni Hills. L. c. vaughani is restricted to the island of Pemba,
and differs from mandanus in having the crown bluish violet rather than green
or greenish blue.
Acknowledgements : For the organising of loans of additional material from
the collections of the East London Museum and the National Museum of
Rhodesia, Bulawayo, I am grateful to Mr. C. D. Quickelberge and Mr.
M. P. Stuart Irwin.
"5
References :
Amadon, D. 1956. Remarks on the Starlings, Family Sturnidae. Amer. Mus. Novit., 1803:
26, 27.
— 1962. Family Sturnidae in the continuation of Peters' Check-list of Birds of the World,
15: 92.
Clancey